GSA Special Publication No.4: Mesozoic and Cainozoic Palynology: Essays in honour of Isabel Cookson

GEOLOGICAL SOCIETY OF AUSTRALIA INCORPORATED

SPECIAL PUBLICATION No. 4

Mesozoic and Cainozoic Palynology: Essays in Honour of Isabel Cookson Editors: J. E. GLOVER and G. PLAYFORD

SYDNEY DECEMBER, 1973

1973 THE GEOLOGICAL SOCIETY OF AUSTRALIA INCORPORATED

Office Bearers 1972-1973 (Elected 23 May 1971)

President: D. A. BROWN Vice-Presidents: A. F. TRENDALL, DOROTHY HILL Hon. Secretary: M. J. RICKARD Hon. Treasurer: K. R. WALKER Hon. Administrative Officer: K. G. MOSHER Hon. Editor: K. A. TOWNLEY Editorial Advisers: R. A. BINNS, K. S. W. CAMPBELL, E. S. HILLS, P. F. HOWARD, R. W. R. RUTLAND, A. F. TRENDALL

GEOLOGICAL SOCIETY OF AUSTRALIA INCORPORATED

SPECIAL PUBLICATION No. 4 Mesozoic and Cainozoic Palynology: Essays in Honour of Isabel Cookson Editors: J. E. GLOVER and G. PLAYFORD

DECEMBER, 1973

1973 THE GEOLOGICAL SOCIETY OF AUSTRALIA INCORPORATED

This volume is published in the Special Publications series of the Geological Society of Australia Reference:—Spec. Pubis geol. Soc. Aust., 4

Printed at The Griffin Press, Adelaide, S.A. Issued December, 1973 Registered in Australia for transmission by post as a book.

Please address inquiries as follows: Editorial Matters: Mr K. A. TOWNLEY, Bureau of Mineral Resources, Geology and Geophysics, P.O. Box 378, Canberra City, A.C.T. 2601. Business Matters: Mr K. G. MOSHER, Hon. Administrative Officer, Geological Society's Office, Gemmology House, 24 Wentworth Avenue, Sydney, N.S.W. 2010. Price to members $Aus8.50 Price to non-members $Aus 12.00 Prices include postage.

DR ISABEL CLIFTON COOKSON

DR ISABEL CLIFTON COOKSON* A Preface to Special Publication No. 4 By GEORGE BAKER More than half-a-century of scientific research work is indeed a meritorious achievement. Dr Isabel Clifton Cookson has joined the small and illustrious group of Australian scientists who can claim this accomplishment. Her studies embrace the disciplines of botany, palaeobotany, and palynology. They have a significant bearing on several aspects of the problems of stratigraphical geology in Australia. Consequently, university personnel and research scientists in Australia and abroad derive much satisfaction from the decision of the Federal Executive of the Geological Society of Australia to publish this symposium-volume in honour of Isabel Cookson. Most of the papers were presented during a two-day symposium on Mesozoic and Cainozoic Palynology, dedicated to Dr Cookson and convened by Dr Geoffrey Playford, at the 43 rd Congress of the Australian and New Zealand Association for the Advancement of Science (Brisbane, May 1971). Isabel Cookson is the doyen of palynologists in Australia, and without doubt is the pioneer of palynology on this continent. In paying tribute to her work, it is hoped that this biographical introduction to the Geological Society's volume in her honour, does adequate justice to one of Australia's well-known women scientists—one who has devoted her life with zeal and energy to the pursuit of knowledge in the field of living and fossil plant micro-organisms. Born in Melbourne on Christmas Day, 1893, Isabel Cookson was educated at the Methodist Ladies' College in the Melbourne suburb of Hawthorn. Here she was a Prefect and a member of the school tennis 'First-Four'. Biology was her favoured subject. Music was also one of her interests. Proficiency as a pianist provided many pleasant moments of relaxation in the years to come. On passing the Junior Public and Junior Commercial Examinations in 1910, and gaining honours in botany and honours in anatomy and physiology in 1911 at the Senior Public Examinations, Isabel Cookson began her

undergraduate studies in 1913 at the University of Melbourne where she concentrated on the biological sciences. The seeds had now been sown for the growth and development of a long life's work of dedication to botanical and palaeobotanical investigations. Nevertheless, her early interest in sport was maintained for a time, and Isabel Cookson represented her Alma Mater in the Women's Tennis 'First-Four' in contests against Adelaide University, held in Adelaide. Obtaining honours in biology in the first two years of her Bachelor of Science degree course, she completed the third and final year with excellent results, gaining Second Class Honours and the Exhibition in Zoology III (December, 1915) and First Class Honours and the Exhibition in Botany III (December, 1915). After graduating B.Sc. in April 1916, majoring in biology, she demonstrated in botany laboratory classes and undertook research work for the Master of Science degree under the guidance of the late Professor A. J. Ewart in the botany section of the old Biology School, University of Melbourne. In later years she conducted tutorial classes in biology in Newman College, University of Melbourne. She had been awarded a Government Research Scholarship in May 1916 for work on the flora of the Northern Territory. This was followed by her winning the McBain Research Scholarship in Biology in December 1916, and a First Class Final Honours Scholarship in Botany in March 1917. She was also recommended for a Government Research Scholarship in April 1917 for the purpose of studying the longevity of cut flowers. Further research work under Professor Ewart, Professor S. M. Wadham (Agriculture Department), and Professor W. H. Lang (University of Manchester, England), and joint research investigations in association with Ethel McLennan (Botany) and the late Frederick Chapman (Commonwealth Palaeontologist in the National Museum of Victoria) in Melbourne, were conducted between 1917 and 1930.

*The Editors regret that Dr Cookson died in Melbourne on 1 July 1973, while this volume was in press.

iv

GEORGE BAKER

In this period, Isabel Cookson was awarded several research grants—one in April 1925 for research in botany, one in 1928 for investigations into crown rot in walnut trees, the research being carried out in the Agriculture Department at Melbourne University. Then in 1929, a further research grant was for work on the Devonian flora in Victoria; another in 1930 was for the purpose of studying fossil plants in the Tertiary sediments at Yallourn, Victoria. In August 1925, a proposal for her to leave for England was approved by Melbourne University, so that she could advance her studies of fungi under Professor Le Rayner in London, and consult on fossil plants in Cambridge with the British palaeobotanical authority Professor Sir A. C. Seward. While in London, she worked on fungi with Ethel McLennan at the Imperial College of Science and Technology. Her research thesis on Early Palaeozoic plants from Alexandra, Victoria, commenced in 1930, led to her gaining the degree of Doctor of Science at Melbourne University, in 1932. The D.Sc. degree was first awarded in Melbourne University in 1887, and first conferred on a woman in 1909; in 1932, Isabel Cookson became the fourth woman graduate to take out this degree in the University of Melbourne. Among her earlier work, investigations of a crown rot of English walnut trees in the Bright district of Victoria revealed fungal hyphae in the diseased areas. In published comments on her paper, Professor S. M. Wadham stated that the description of the fungus, and its occurrence and life history, brought out important facts of particular significance to the cultivation of walnuts. Turning from the purely botanical aspect of her early research work, we find she also scored a major triumph on the palaeobotanical side by recording and describing some of the earliest known fossil land plants of vascular character. These were from Walhalla, Victoria and from the Rhyll bore, Victoria, and her first paper on the subject was published in 1926. In May 1926, Isabel Cookson journeyed to England a second time. On this occasion, she acted as mycologist in cotton research at the University of Manchester until February 1927. This provided an additional opportunity for collaboration with Professor Lang in studying the earliest known vascular land plants.

Returning to Australia for a period of five years, she then obtained a Grisedale Research Scholarship for 1933-1934 for still further investigation into the nature and significance of the early land plants in Victorian older Palaeozoic sediments. Three important papers arose from Isabel Cookson's research work carried out in association with Professor Lang in the Botany Department at the University of Manchester in 1926 and 1933. These were published in 1927, 1930, and 1935 in co-authorship with Professor Lang. Thus it became well established that some of the earliest known vascular land plants on Earth occurred in the older Palaeozoic sediments of Victoria. Much of the field sampling at Walhalla and other Victorian localities was carried out by Isabel Cookson herself, at a period, and in country, where the going was arduous to say the least. Use was also made of specimens collected by officers of the Geological Survey of Victoria. Worthy of especial note is the collection of certain specimens from along the Yarra Track which revealed examples of the Siluro-Devonian genus Monograptus exposed on the same bedding plane as genera of the vascular land plants—I understand that credit for the discovery of some of these important specimens (now dated as Early Devonian) goes to Isabel Cookson. As an outcome of this work, W. N. Croft and W. H. Lang honoured Isabel Cookson in their paper of 1942 {Phil. Trans. R. Soc. Lond., (Ser. B), 231, p. 131) by creating the new genus Cooksonia for dichotomously branched, slender leafless axes of plants terminating in large sporangia, found in Lower Devonian sediments in Wales. Lecturing duties commenced for Isabel Cookson in 1930, when a separate Botany Department had become established in the grounds of the University of Melbourne. The first evening course in first year Botany was inaugurated under her charge in this year. As a member of this class—the first series of evening Botany classes in Melbourne University—I well remember the lucid character of her lectures and her thoroughly competent handling of a most interesting series of laboratory classes, which, being conducted at nighttime, necessitated the examination of all microscopic preparations under artificial illumination. Her stimulating approach to teaching and her patient attention to detail were most encouraging to newcomers to the intricacies of

DR. ISABEL CLIFTON COOKSON the science of botany—and there was a high proportion of newcomers to botany in that first evening course. For Isabel Cookson, lecturing and demonstrating in first year Botany to evening students continued until 1947, and courses in p a l e o botany were also delivered to senior students of botany. At this stage of her career, a little over thirty years of University teaching and research work had elapsed. The following quarter-of-a-century was to become equally as active and stimulating for her in the field of palaeobotanical research. For 1948-1949, she was awarded a Leverhulme Research Grant in the University of Manchester. During the tenure of this grant, she prepared a paper on Yeringian (Lower Devonian) plant remains from Lilydale, Victoria in consultation with Professor W. H. Lang, F.R.S. at the Manchester Museum. The results of this work were published in the 1949 Memoirs of the National Museum of Victoria, Melbourne. Then, in 1949, Isabel Cookson was appointed Leader of the Pollen Research Unit in Melbourne University. The establishment of a unit of this nature had been mooted and discussed in 1947, with the concept that it would be advantageous, and certainly most useful, to make detailed palaeobotanical studies, with emphasis on pollen analysis, of Victoria's vast brown coal and associated deposits. Sponsored by C.S.I.R.O., and with annual contributions from the State Electricity Commission of Victoria, it was agreed with the University of Melbourne that the headquarters of the Pollen Unit be located in their Botany Department. It was under the administration of Professor J. S. Turner, Head of the Botany Department, and Dr A. B. Edwards of the C.S.I.R.O. Mineragraphic Investigations Section which was housed in the University of Melbourne. Leading up to her appointment to the Pollen Research Unit, the only one of its kind in Australia, Isabel Cookson had spent sabbatical leave in 1948 at the Palynological Laboratory, Bromma (Stockholm), Sweden, backed by a C.S.I.R.O. Research Grant. This followed shortly upon her completion and publication of investigations of plant microfossils in the lignites from the Kerguelen Archipelago. In Stockholm, she reaped the benefit of consultations with Dr G. Erdtman the world authority on pollen grains and microspores.

v

This helped to pave the way for the outstanding success of the Pollen Unit during its active and fruitful existence of the following seventeen years or so. From the date of its inception in 1949, some twenty-five to thirty papers were published by members of the Unit on pollen grains, spores, and macrofossils of plants in the brown coal and other Tertiary sediments of Victoria. Isabel Cookson was sole author of over fifty per cent of these papers, joint author in the remainder with her coworkers, Suzanne L. Duigan, Kathleen M. Pike, and Mary E. Dettmann. If one thumbs through the 539 pages of that excellent volume 'Pollen Morphology and Plant Taxonomy—Angiosperms (An Introduction to Palynology I ) ' by G. Erdtman, published by the Chronica Botanica Co., in Waltham, Massachusetts, U.S.A. in 1952, one is immediately struck by the fact that the only name of a joint author given for any one of the many families of the angiosperms dealt with, is that of Isabel Cookson. The large family Proteaceae was covered jointly by Isabel Cookson and G. Erdtman on pages 339-369, with Figures 198-214. There are no joint authors for any of the other families. Assuredly this is a great tribute to Australia's premier pollen expert, for, as Mr. George Baragwanath of the State Electricity Commission of Victoria recently remarked, Erdtman's book is virtually the research workers' 'bible' in palynology. The work for this joint authorship was conducted during Isabel Cookson's visit to the Stockholm Palynological Laboratory in 1948. In 1952, she was classified as a Research Fellow in the Botany Department of the University of Melbourne, with the status of Senior Lecturer. Once the science of palynology was wellestablished in Australia by Isabel Cookson, its pioneer turned her attentions more particularly to other groups of plant-like microfossils that had come under her notice from time to time. During the painstaking separation techniques necessary for isolating pollen grains, microspores, and megaspores from their various sedimentary host rocks, including carbonaceous sandstones, siltstones, and argillites, certain micro-organic remains had been observed which she subsequently proved to be representatives of such microplanktonic groups as the dinoflagellates and the hystrichosphaerids.

vi

GEORGE BAKER

Although her first paper relating to the hystrichosphaerids appeared in 1953, it is worthy of note that Isabel Cookson had observed these microfossils as early as 1945. They were in the first preparations we had made in the Geology Department, University of Melbourne from the Nelson bore sediments, expressly for determination of the pollen grain and spore content by Isabel Cookson. Identification and description of the various forms present, however, had to remain in abeyance pending progress of the systematic examination of pollen grains and spores from a number of living and fossil plants in Australia and New Guinea. Faced with the challenge of elucidating the nature, occurrence, and significance of fossil microplankton in Australian Tertiary and Mesozoic sediments, the indefatigable Isabel Cookson spent 1954 sabbatical leave in France. In Paris, she worked with Professor G. Deflandre on those intriguing unicellular marine organisms of microscopic size, of obscure origin, and of uncertain affinities—the dinoflagellates and the hystrichosphaerids. This work immediately paid dividends, for in 1955, it was shown for the first time, from studies of the microplankton content, that Cretaceous sediments of marine origin occurred in Victoria—in the lower portions (5,782—6,192 ft. i.e. 1757—1882 m) of the Nelson bore, western Victoria. These and related sediments subsequently became a target for boring operations during the earlier phases of the recently stepped-up search for oil in Victoria. In the following years, a number of oil exploration companies and geological surveys called on Isabel Cookson's knowledge of fossil pollen grains, microspores, megaspores, and organic-walled microplankton, in helping to elucidate and correlate the sedimentary sequences in their exploratory boreholes. Once again she had pioneered a new branch of scientific research in Australia—the isolation and description of fossil microplankton in our Tertiary and Mesozoic sedimentary rocks. This has proved to be of particular significance and value in studies of bore core samples, where the amount of material recovered is limited, and especially where other types of fossils or diagnostic species are wanting. From November 1956 to April 1957, Isabel Cookson again visited Europe. The intention was to work in France to advance her studies of the microplankton, but this plan did not

eventuate in all its details. Consequently she journeyed on to Reutlingen in Germany for consultation and collaboration with Professor A. Eisenack in the University of Tubingen. Thus began an association which resulted in the publication of many joint papers dealing with microplankton in the Tertiary and Mesozoic sediments of Australia and New Guinea. Her last overseas leave as a staff member of the University of Melbourne, was from October 1958 to March 1959, when she again worked with Professor Eisenack in Reutlingen and later with Svein Manum in Oslo, Norway. With the effluxion of time, official retirement from the Melbourne University staff came about in 1959. This, however, was by no means the end of Isabel Cookson's research career, for research was her life, and to research she tenaciously adhered. Now, some twelve years later, she is still writing papers on Australian fossil microplankton. Twenty-seven research papers have been published, principally with co-authors, since her official retirement; another one is going through the press at this stage, and others are planned for the immediate future. Post-retirement research investigations have been possible for her through the generosity of the University of Melbourne Botany Department in providing room and laboratory space, and in supplying the necessary equipment and reagents. In one way or another, she has now been associated with her Alma Mater for the past fifty-eight years—truly a remarkable connection. Her interest in active research has also been maintained by relatively frequent visits overseas in the last ten years, for collaboration with fellow workers on the latest developments in microplankton studies. These visits included Germany and Great Britain in 1961, the U.S.A. and Great Britain in 1963, Germany in 1965 and again in 1968 and 1970. All were financed personally, mostly for the specific purpose of furthering publication of collaborative research work with overseas scientists who showed a deep interest in the Australian fossil microplankton. The fruitfulness of these visits and her unending capacity for research work is revealed by reference to published papers of the post-1959 period set out in the list accompanying this biography. These were produced jointly with Professor Eisenack (16 papers), S. Manum (3), N. F. Hughes (1), and Lucy M. Cranwell (1) among overseas workers; and with Mary E.

vii

DR. ISABEL CLIFTON COOKSON

Dettmann (1), B. E. Balme (1), and W. K. Harris (1) among Australian workers. Isabel Cookson was elected to Associate Membership of the Royal Society of Victoria in 1916, the year of her B.Sc. graduation. She transferred to full Membership in 1959, and for the past twelve years has been a Life Member of the Society. During fifty-five years membership, she has contributed twenty-eight papers to this Society's Proceedings. From 1959 to 1962, she acted as an Honorary Associate in Palaeontology to the National Museum of Victoria in Melbourne, and contributed two papers to the Museum's Memoirs. It is in the extensive collections of this institution that many of her microscopic preparations of plant microfossils from the eastern States of Australia have been lodged, more especially her types and figured specimens. Those prepared from Western Australian sediments she has lodged for permanent keeping in the collections of the Geological Survey of Western Australia, more especially holotypes and hypotypes. Among the honours that came her way, Isabel Cookson was invited to deliver a lecture at the official opening of the Birbal Sahni Institute for Palaeobotany at Lucknow, India. At this ceremony, she was appointed official Australian representative of the Australian National Research Council. The Botanical Society of America elected her a Corresponding Member in 1957, for distinguished services to botany. Isabel Cookson can look back along the past fifty and more years of her scientific

career, secure in the knowledge that she has come up with several firsts in the domain of original research work in palaeobotany in Australia, confident that her work has been of significant value to more than one scientific discipline, and contented in the thought that her many collaborators in different realms of botanical and palaeobotanical research investigations have reaped mutual benefits from associating in the work. To have collaborated with seventeen other scientists in the preparation for publication of fifty joint papers, is truly a splendid example of active co-operation in research. She has published thirty-five papers as sole author, and the eighty-five papers with which she has been concerned as joint or as sole author, have been published in twenty-six different journals, thirty one per cent of them overseas. Her colleagues warmly appreciate her diligence and contributions to botanical and palaeobotanical research. As mentioned previously, this volume represents the published outcome of the 1971 A.N.Z.A.A.S. Symposium on Mesozoic and Cainozoic Palynology, held in commemoration of Dr Cookson and her outstanding contributions in that field of scientific endeavour. The success of that Symposium was assured by the excellent response from a score of contributors working in the universities, in geological surveys, and in oil-exploration companies, on an Australia-wide basis. Surely this is eloquent testimony to the esteem and high regard held for the pioneer of palynology in Australia— Isabel C. Cookson.

BIBLIOGRAPHY OF ISABEL C. COOKSON Papers are listed in chronological order. 1921: Floral abnormalities in the genera Eriostemon and Glossodia. Proc. R. Soc. Vict., 33, pp. 32-38.

COOKSON, ISABEL C . ,

MCLENNAN, ETHEL, & COOKSON, ISABEL C . ,

1923:

Additions to the Australian Ascomycetes, No. 1. Proc. R. Soc. Vict., 35, pp. 153-158. COOKSON, ISABEL C., 1926: On the occurrence of the Devonian genus Arthrostigma in Victoria. Proc. R. Soc. Vict., 38, pp. 65-68. CHAPMAN, F . , & COOKSON, ISABEL C . ,

1926:

A

revision of the 'Sweet' collection of Triassic plant remains from Leigh Creek, South Australia. Trans. R. Soc. S. Aust., 50, pp. 163178. MCLENNAN, ETHEL, & COOKSON, ISABEL C . ,

1926:

Additions to Australian Ascomycetes, No. 2. Proc. R. Soc. Vict., 38, pp. 69-76.

LANG, W . H . , & COOKSON, ISABEL C . ,

1927:

On

some early Palaeozoic plants from Victoria, Australia. Mem. Proc. Manchr lit. phil. Soc., 71, p p . 4 1 - 5 1 . ISABEL C . , 1928: The structure and development of the perithecium of Melanospora zamiae Corda. Ann. Bot., 42, pp. 255269. , 1929: An account of a crown rot of English walnut trees in Victoria. Proc. R. Soc. Vict., 42, pp. 5-25. LANG, W. H., & COOKSON, ISABEL C . , 1930: Some fossil plants of early Devonian type from the Walhalla Series, Victoria, Australia. Phil. Trans. R. Soc. Lond., (Ser. B), 219, pp. 133-163. COOKSON,

GEORGE BAKER

viii

, , 1935: On a flora, including vascular land plants, associated with Monograptus, in rocks of Silurian age, from Victoria, Australia. Phil. Trans. R. Soc. Lond., (Ser. B), 224, pp. 421-449. COOKSON, ISABEL C . , 1 9 3 5 : On plant remains from the Silurian of Victoria, Australia, that extend and connect floras hitherto described. Phil. Trans. R. Soc. Lond., (Ser. B), 225, pp. 127-148.

, 1937: The occurrence of fossil plants at Warrentinna, Tasmania. Pap. Proc. R. Soc. Tasm. pp. 73-77. , 1937: On Saprolegnia terrestris sp. nov., with some preliminary observations on Victorian Saprolegniales. Proc. R. Soc. Vict., 49, pp. 235-242. , 1937: Fossil wood from Upper Devonian rocks at Mansfield, Victoria. Proc. R. Soc. Vict., 50, pp. 182-189. , 1945: Pollen content of Tertiary deposits. Aust. J. Sci., 7, pp. 149-150. , 1945: Records of plant remains from the Upper Silurian and Early Devonian rocks of Victoria. Proc. R. Soc. Vict., 56, pp. 119-122. , 1946: Pollens of Nothofagus Blume from Tertiary deposits in Australia. Proc. Linn. Soc. N.S.W., 71, pp. 49-63. , 1947: Note on the pollen of Nothofagus gunnii (Hook.) Oerst. Proc. R. Soc. Vict., 58, pp. 1-2. , 1947: On fossil leaves (Oleaceae) and a new type of fossil pollen grain from Australian brown coal deposits. Proc. Linn. Soc. N.S.W., 72, pp. 183-197. , 1947: Fossil fungi from Tertiary deposits in the Southern Hemisphere. Pt I. Proc. Linn. Soc. N.S.W., 72, pp. 207-214. , 1947: Plant microfossils from the lignites of Kerguelen Archipelago. B.A.N.Z. Antarct. Res. Exped. 1929-31, Reports, Series A, 2, pp. 127-142. , 1949: Yeringian (Lower Devonian) plant remains from Lilydale, Victoria, with notes on a collection from a new locality in the Siluro-Devonian sequence. Mem. natn. Mus., Vict., 16, pp. 117-131. , 1950: Fossil pollen grains of proteaceous type from Tertiary deposits in Australia. Aust. /. Sci. Res., Ser. B, 3, pp 166-177. , & DUIGAN, SUZANNE L., 1 9 5 0 : Fossil Banksieae from Yallourn, Victoria, with notes on the morphology and anatomy of living species. Aust. J. Sci. Res., Ser. B, 3, pp. 133-165. 1951: Tertiary Araucariaceae y from south-eastern Australia, with notes on living species. Aust. J. Sci. Res., Ser. B, 4, pp. 415-449. COOKSON, ISABEL C . , 1952: Identification of Tertiary pollen grains with those of New Guinea and New Caledonian beeches. Nature, Lond., 170, p. 127.

, & ERDTMAN, G.,

1952:

Proteaceae;

in

Erdtman, G., Pollen Morphology and Plant Taxonomy—Angiosperms (An Introduction to Palynology / ) , pp. 339-369. Chronica Botanica Co., Waltham, Mass. COOKSON, ISABEL C . , 1953: The identification of the sporomorph Phyllocladidites with Dacrydium and its distribution in southern Tertiary deposits. Aust. J. Bot., 1, pp. 64-70. , 1953: Records of the occurrence of Botryococcus braunii, Pediastrum and the Hystrichosphaerideae in Cainozoic deposits of Australia. Mem. natn. Mus., Vict., 18, pp. 107-123. ,1953: On Macrozamia hopeites—an early Tertiary cycad from Australia. Phytomorphology, 3, pp. 306-312. , 1953: Difference in microspore composition of some samples from a bore at Comaum, South Australia. Aust. J. Bot., 1, pp. 462-473. , & PIKE, KATHLEEN M., 1 9 5 3 : A contribution to the Tertiary occurrence of the genus Dacrydium in the Australian region. Aust. J. Bot., 1, pp. 474-484. , , 1953: The Tertiary occurrence and distribution of Podocarpus (section Daery carpus) in Australia and Tasmania. Aust. J. Bot., 1, pp. 71-82. CLIFFORD, H .

T . , & COOKSON, ISABEL C . ,

1953:

Muscites yallournensis, a fossil moss capsule from Yallourn, Victoria. Bryologist, 56, pp. 53-55. COUPER, R . A . , HARRIS, W . F . , COOKSON, ISABEL C . , & DUIGAN, SUZANNE L., 1953: Termi-

nology and nomenclature for fossil pollens and spores. Proc. Int. Bot. Congr7, pp. 890-891. COOKSON, ISABEL C . , 1954: The Cainozoic occurrence of Acacia in Australia. Aust. J. Bot., 2, pp. 52-59. , 1954: The occurrence of an older Tertiary microflora in Western Australia. Aust. J. Sci., 17, pp. 37-38. , 1954: A palynological examination of No. 1 bore, Birregurra, Victoria. Proc. R. Soc. Vict., 66, pp. 119-128. , 1954: Recent additions to our knowledge of Australian Tertiary floras. Rapp. Commun., Int. Bot. Congr., 8, pp. 223-227. , & PIKE, KATHLEEN M., 1954: The fossil occurrence of Phyllocladus and two other podocarpaceous types in Australia. Aust. J. Bot., 2, pp. 60-68. , , 1954: Some dicotyledonous pollen types from Cainozoic deposits in the Australian region. Aust. J. Bot., 2, pp. 197-219. COOKSON, ISABEL C . , & SINGLETON, O . P . ,

1954:

The preparation of translucent fossils by treatment with hydrofluoric acid. News Bull, geol. Soc. Aust., 2, pp. 1-2.

DR. ISABEL CLIFTON COOKSON DEFLANDRE,

G.,

& COOKSON,

ISABEL

C.,

1954:

Sur le microplancton fossile conserve dans diverse roches sedimentaires australiennes s'etageant du Cretace inferieur au Miocene superieur. C.R. Acad. Sci., Paris, 239, pp. 1235-1238. ISABEL C., 1 9 5 5 : The occurrence of Palaeozoic microspores in Australian Upper Cretaceous and Lower Tertiary sediments. Aust. J. Sci., 18, pp. 5 6 - 5 8 . , & PIKE, KATHLEEN M . , 1 9 5 5 : The pollen morphology of Nothofagus BL. sub-section Bipartitae Steen. Aust. J. Bot., 3, pp. 197-

COOKSON,

206. BAKER, G . , & COOKSON, ISABEL C . , 1 9 5 5 : A g e

of

Nelson bore sediments. Aust. J. Sci., 17, pp. 133-134. DEFLANDRE,

G.,

& COOKSON,

ISABEL

C.,

1955:

Fossil microplankton from Australian Late Mesozoic and Tertiary sediments. Aust. J. mar. Freshw. Res., 6, pp. 242-313. COOKSON, ISABEL C . , 1956: Pollen grains of the Ephedra type in Australian Tertiary deposits. Nature, Lond., 177, pp. 47-48. , 1956: Additional microplankton from Australian Late Mesozoic and Tertiary sediments. Aust. J. mar. Freshw. Res., 7, pp. 183-191. , 1957: On some Australian Tertiary spores and pollen grains that extend the geological and geographical distribution of living genera. Proc. R. Soc. Vict., 69, pp. 41-53. DUIGAN,

SUZANNE

L.,

&

COOKSON,

ISABEL

C.,

1957: The occurrence of Azolla filiculoides L. and associated vascular plants in a Quaternary deposit in Melbourne, Australia. Proc. R. Soc. Vict., 69, pp. 5-13. COOKSON,

ISABEL

C.,

&

DETTMANN,

MARY

E.,

1958: Cretaceous 'megaspores' and a closely associated microspore from the Australian region. Micropaleontology, 4, pp. 39-49. , , 1958: Some trilete spores from Upper Mesozoic deposits in the eastern Australian region. Proc. R. Soc. Vict., 70, pp. 95-128. COOKSON, ISABEL C . , & EISENACK, A., 1 9 5 8 :

Microplankton from Australian and New Guinea Upper Mesozoic sediments. Proc. R. Soc. Vict., 70, pp. 1 9 - 7 9 . COOKSON, ISABEL C . , 1959: Fossil pollen grains of Nothofagus from Australia. Proc. R. Soc. Vict., 71, pp. 25-30. , & DETTMANN, MARY E . , 1 9 5 9 : Microfloras in bore cores from Alberton West, Victoria. Proc. R. Soc. Vict., 71, pp. 31-38. , , 1959: On Schizosporis, a new form genus from Australian Cretaceous deposits. Micropaleontology, 5, pp. 213-216. , , 1959: Cyclosporites, Cretaceous microspore: corrected name. Aust. J. Sci., 21, p. 260.

COOKSON,

ISABEL

ix C.,

&

EISENACK,

A.,

1960:

Microplankton from Australian Cretaceous sediments. Micropaleontology, 6, pp. 1-18. , , 1960: Upper Mesozoic microplankton from Australia and New Guinea. Palaeontology, 2, pp. 2 4 3 - 2 6 1 . COOKSON,

ISABEL C . ,

& MANUM,

S.,

1960:

On

Crassosphaera, a new genus of microfossils from Mesozoic and Tertiary deposits. Nytt Mag. Bot., 8, pp. 5 - 1 0 . EISENACK, A., & COOKSON, ISABEL C., 1 9 6 0 : Microplankton from Australian Cretaceous sediments. Proc. R. Soc. Vict., 72, pp. 1-11. COOKSON, ISABEL C . , 1 9 6 1 : Hoegisporis, a new Australian Cretaceous form genus. Palaeontology, 3, pp. 4 8 5 - 4 8 6 . , & DETTMANN, MARY E., 1 9 6 1 : Reappraisal of the Mesozoic microspore genus Aequitriradites. Palaeontology, 4, pp. 4 2 5 - 4 2 7 . COOKSON, ISABEL C . , & EISENACK, A . , 1 9 6 1 :

Ter-

tiary microplankton from the Rottnest Island bore, Western Australia. /. Proc. R. Soc. West. Aust., 44, pp. 3 9 - 4 7 . , , 1961: Upper Cretaceous microplankton from the Belfast No. 4 bore, southwestern Victoria. Proc. R. Soc. Vict., 74, pp. 69-76. COOKSON,

ISABEL

C.,

&

BALME,

B.

E.,

1962:

Amosopollis cruciformis gen. et sp. nov., a pollen tetrad from the Cretaceous of Western Australia. J. Proc. R. Soc. West. Aust., 45, pp.

97-99.

COOKSON,

ISABEL

C.,

&

EISENACK,

A.,

1962:

Additional microplankton from Australian Cretaceous sediments. Micropaleontology, 8, pp.

485-507.

, , 1962: Some Cretaceous and Tertiary microfossils from Western Australia. Proc. R. Soc. Vict., 75, pp. 2 6 9 - 2 7 3 . COOKSON,

ISABEL

C.,

& HUGHES,

N.

F.,

1964:

Microplankton from the Cambridge Greensand (mid-Cretaceous). Palaeontology, 7, pp. 37-59. COOKSON,

ISABEL

C.,

& MANUM,

S.,

1964:

On

Deflandrea victoriensis n. sp., D. tripartita Cookson and Eisenack and related species. Proc. R. Soc. Vict., 77, pp. 5 2 1 - 5 2 4 . MANUM, S., & COOKSON, ISABEL C . ,

1964:

Cre-

taceous microplankton in a sample from Graham Island, Arctic Canada, collected during the Second 'Fram' Expedition ( 1 8 9 8 1 9 0 2 ) , with notes on microplankton from the Hassel Formation, Ellef Ringnes Island. Skr. norske Vidensk-Akad. Mat.-Naturv Kl., 17, pp. 1 - 3 6 . COOKSON, ISABEL C . , 1 9 6 5 : On a new species of Hoegisporis Cookson. Palaeontology, 8, pp. 39-40.

, 1965: Cretaceous and Tertiary microplankton from south-eastern Australia. Proc. R. Soc. Vict., 78, pp. 8 5 - 9 3 .

GEORGE BAKER , 1965: Microplankton from the Paleocene COOKSON, ISABEL C., & EISENACK, A., 1967: Some Early Tertiary microplankton and pollen Pebble Point Formation, south-western Vicgrains from a deposit near Strahan, western toria. Part 1. Proc. R. Soc. Vict., 78, pp. Tasmania. Proc. R. Soc. Vict., 80, pp. 131137-141. 140. , & EISENACK, A., 1965: Microplankton , , 1967: Some microplankton from from the Browns Creek Clays, S.W. Victoria. the Paleocene Rivernook Bed, Victoria. Proc. Proc. R. Soc. Vict., 79, pp. 119-131. R. Soc. Vict., 80, pp. 247-258. , 1965: Microplankton from the , , 1968: Microplankton from two Dartmoor Formation, S.W. Victoria. Proc. R. samples from Gingin Brook No. 4 borehole, Soc. Vict., 79, pp. 133-137. Western Australia. J. Proc. R. Soc. West. , , 1965: Microplankton from the Aust., 51, pp. 110-122. Paleocene Pebble Point Formation, south, , 1969: Some microplankton from western Victoria. Part 2. Proc. R. Soc. Vict., two bores at Balcatta, Western Australia. J. 79, pp. 139-146. Proc. R. Soc. West. Aust., 52, pp. 3-8. HARRIS, W . K . , & COOKSON, ISABEL C . , 1 9 6 5 : T h e , , 1970: Die Familie der Lecanielstratigraphy of the Comaum No. 2 bore—A laceae n. fam.—Fossile Chlorophyta, Volreinterpretation. Aust. J. Sci., 28, pp. 25-26. vocales? Neues Jb. Geol. Paldont. Mh., 1970 (6), pp. 321-325. COOKSON, ISABEL C . , & CRANWELL, L U C Y M . , 1970: Cretaceous microplankton 1967: Lower Tertiary microplankton, spores from the Eucla Basin, Western Australia. and pollen grains from southernmost Chile. Proc. R. Soc. Vict., 83, pp. 137-158. Micropaleontology, 13, pp. 204-216.

X

5

5

George Baker, 146 Wimbledon Avenue, Mt Eliza, Victoria 3930.

CONTENTS STRATIGRAPHIC, PALAEOECOLOGICAL, AND PALAEOBOTANICAL ASPECTS OF SPORE-POLLEN FLORAS MARY E .

DETTMANN:

Angiospermous Pollen from Albian to Turonian Eastern Australia HELENE A.

Sediments

of

3

MARTIN:

Upper Tertiary Palynology in Southern New South Wales

-

-

35

Upper Cretaceous-Eocene Spore-Pollen Zonation, Offshore Gippsland Basin, Australia -

55

L E W I S E . STOVER & P . RICHARD E V A N S :

A. R. H.

MARTIN:

Reappraisal of Some Palynomorphs of Supposed Proteaceous Affinity. 1. The Genus Beaupreaidites Cookson ex Couper and the Species Proteacidites hakeoides Couper . D.

M.

D.

BURGER:

The Ecological Significance of Tropical Mangroves in the Early Tertiary Floras of Southern Australia Spore Zonation and Sedimentary History of the Neocomian, Great Artesian Basin, Queensland J. G .

79

87

DOUGLAS:

Spore-Plant Relationships in Victorian Mesozoic Cryptogams N.

73

CHURCHILL:

-

119

J . DE JERSEY:

Rimulate Pollen Grains from the Lower Mesozoic of Queensland ROBIN

HELBY:

Review of Late Permian and Triassic Palynology of New South Wales

127 141

STUDIES OF NON-CALCAREOUS MICROPLANKTON WAYNE K .

HARRIS:

Tertiary Non-marine Dinoflagellate Cyst Assemblages from Australia LEWIS E .

Palaeocene and Eocene Species of Deflandrea Victorian Coastal and Offshore Basins, Australia

in -

167

The Numerical Analysis of Modern Pollen Spectra from Northeast Queensland Rain-forests

191

STATISTICAL A. P.

J.

159

STOVER:

APPROACHES

TO

POLLEN

(Dinophyceae) -

ANALYSIS

KERSHAW:

GUPPY,

P.

MILNE,

M.

GLIKSON, & M .

MOORE:

Further Developments in Computer Assistance to Pollen Identification

201

INDEXES Index to Described and/or Illustrated Species Author Index

-

207 209

STATIGRAPHIC, PALAEOECOLOGICAL, AND PALAEOBOTANICAL ASPECTS OF SPORE-POLLEN FLORAS

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN SEDIMENTS OF EASTERN AUSTRALIA By MARY E. DETTMANN (With 1 Table, 3 Text-Figures and 6 Plates) ABSTRACT

Plant microfossils having characters consistent with angiospermous pollen morphology are minor constituents of Albian to Turonian palynological floras in the eastern Australian region. Within the area studied, from Bathurst Island in the north to the Otway Basin in the south, the earliest occurring types (of late middle-late Albian age) are tricolpate to tricolpoidate, reticulate species. These diversified and were joined in late Albian to early Cenomanian strata by monosulcate forms. By middle-late Cenomanian times trichotomosulcate and tricolporate species became established and during the Turonian triporate and polyporate forms were introduced. This sequential arrangement diverges somewhat from those that have been recorded from the Americas, Europe, and west Africa; for instance, the first Australian occurrences of the monosulcate form genus Clavatipollenites (late Albian) postdates the appearance of tricolpates, whereas from North America and Europe the genus has been shown to precede the inception of tricolpates and to occur in strata as old as the Barremian. Composition of the Australian Albian and Cenomanian angiospermous pollen suites is relatively uniform throughout the region; some latitudinal differentiation is evident in the later (Turonian) florules. Twenty angiospermous pollen species are described and illustrated by means of the light and scanning electron microscopes. Two new genera, Phimopollenites and Senectotetradites, are proposed, and are typified by P. pannosus (Dettmann & Playford) comb. nov. and S. varireticulatus sp. nov. respectively. In addition the following species are newly instituted herein: Tricolpites cooksonae, Nyssapollenites squamosus, N. lanosus, Triorites punctulatus, and Senectotetradites fistulosus.

INTRODUCTION contain angiosperm elements bearing close In the preface to this volume it has been affiliations with the present-day indigenous noted that much of Dr Isabel Cookson's flowering plants of Australia. Nevertheless, the earlier palynological work was devoted to the time and place of origin and the pre-Tertiary study of angiospermous pollen grains from phylogeny of the extant angiosperm groups the Australian region. These were described have remained virtually unknown. The present and illustrated by Dr Cookson and her col- investigation was undertaken with these problaborators from Early Tertiary to Recent lems in mind and the results, tentative though floras, and important data accrued thereby on they may be in some respects, are here offered the evolution and migration of the Australian as a tribute to Dr Cookson. angiosperm flora which, despite its endemic The current work is a documentation and nature, has distinct affiliations with past and discussion of angiospermous pollen extracted present-day floras of Africa, Antarctica, New from Albian, Cenomanian, and Turonian Zealand, and South America (Burbidge, strata within the Otway and Great Artesian 1960; Takhatajan, 1969). Basins and on Bathurst and Melville Islands. Dr Cookson's engaging enthusiasm, energy, Previous studies of Australian Cretaceous and encouragement led to my own interest in angiospermous pollen grains were carried out palynology, and through an initial collabora- by Dettmann & Playford (1968, 1969) and tion with her (Cookson & Dettmann, 1959) Burger (1971). In their 1968 paper, Dettmann I gained some appreciation of Lower Ter- & Playford described and illustrated several tiary angiospermous floras of eastern Victoria. species of tricolpate, triporate, and polyporate That investigation together with prior and sub- forms which were used subsequently (Dettsequent works initiated by Dr Cookson demon- mann & Playford, 1969) for biostratigraphic strated that Australian Tertiary pollen floras zonation of Albian and Upper Cretaceous Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

4

MARY E. DETTMANN

Fig. 1. Areas of Cretaceous sedimentation in eastern Australia.

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 5 sequences in southeastern Australia. Dettmann here to be more likely of gymnospermous & Playford (1969) also demonstrated that affinity. The natural alliance of Amosopollis within Australia, the earliest angiospermous is less certain, but an algal derivation seems grains occur in late Albian sediments and that probable. by Senonian times Nothofagus Bl. and the Proteaceae had become established. Burger MATERIAL (1971) described tricolpate angiospermous The samples from which the angiospermous pollen from Albian-?early Cenomanian horizons of the Eromanga Basin, Queensland. pollen described in this paper were obtained From associated marine faunas Burger was are from Cretaceous sequences developed on able to date precisely (as late middle Albian) Bathurst and Melville Islands and in the Great the inception of undoubted angiospermous Artesian and Otway Basins (Figs 1, 2). The sediments include both marine and nonpollen in his area of study. marine horizons of Albian, Cenomanian, and Biostratigraphic zonation of latest Creta- Turonian Precise datings are based priceous strata of the Gippsland Basin proposed marily on age. molluscan, foraminiferal, and dinoby Stover & Evans (1973) is a refinement of flagellate evidence from marine strata. KnowDettmann & Playford's (1969) zonation of the spore-pollen content of the scheme for that interval and is based, at least ledge fossiliferous marine strata enables recognition in part, upon angiosperm pollen taxa which of the stages, in varying Stover & Evans describe and illustrate. The in non-marine sections. degrees of precision, latter authors also demonstrate a palynological pertaining to the samples used in method of distinguishing between latest Cre- thisDetails study and to the sections from which they taceous and earliest Tertiary sediments within were collected are given below. southeastern Australia. Two Cretaceous genera thought to bear Bathurst and Melville Islands some resemblance to angiospermous grains Bathurst and Melville Island samples were described by Cookson (1961) and Cook- areThe from outcropping marine sequences which son & Balme (1962) as Hoegisporis and contain well-preserved ammonite faunas dated Amosopollis. Hoegisporis has distally dismiddle or late Cenomanian and Turonian placed peripherial thickenings and an area of as(Wright, 1963; Skwarko, 1966). Albian amthin exine over the ?distal pole; it is considered monites have been identified in a sequence between Ticklitipinapitti and Piplyanyamili Creek, Bathurst Island (Wright, 1963; pp. GREAT A R T E S I A N 604-5, 612; Fig. 3A) but these are thought BASIN BATHI to have been derived. OTWAY The study of dinoflagellate assemblages MELV ' I L L E BASIN MORNINGTON OODNADATTA obtained from the sediments provides inde\SLfikNDS ISLAND pendent and confirmatory evidence for Cenomanian and Turonian ages. Of particular interest is the occurrence of Litosphaeridium I1I11 siphoniphorum (Cookson & Eisenack) Davey TURONIAN & Williams, 1966 which in England and France 1 is confined to, and a useful index species for, the Cenomanian (Clarke & Verdier, 1967; CENOMANIAN 1 1 Davey, 1970); the species appears to be simi1 larly restricted on Bathurst and Melville Islands. Elsewhere in Australia L. siphoni1 1 UPPER phorum is widely distributed in Western Australian sedimentary basins (Cookson & MIDDLE Eisenack, 1958, 1968) and in the northern portion of the Great Artesian Basin (present LOWER study). Precise ages of some of these sediAPTIAN ments are in doubt, but some at least are Cenomanian. It is possible, therefore, that L. siphoniphorum may prove to be a useful CenoFig. 2. Temporal relationships of sediments manian index in Australia as elsewhere. studied. ALBIAN

1

Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

6

MARY E. DETTMANN

Bathurst Island Turonian sediments contain the first appearances of Deflandrea acuminata Cookson & Eisenack, 1958 and lack L. siphoniphorum. A similar situation exists in the Carnarvon Basin, Western Australia, where D. acuminata (s. str.) occurs in beds which are broadly dated as Cenomanian-early Turonian, and succeed those containing L. siphoniphorum (Cookson & Eisenack, 1958). D. acuminata is known also from the Otway Basin (Douglas, 1961; Evans, 1966; present study); some, if not all, of these occurences are in Turonian strata. The following samples are the source of figured angiospermous grains from Bathurst and Melville Islands: 1. Bathurst Island. The samples were collected by Shell Development (Australia) Pty Ltd from southern coastal sections (see also Wright, 1963, Fig. 1). (a) Toyungimpi, sample Ni 36, 2.5 ft (0.75 m) above Tapara bed; Cenomanian. (b) Poupanderie, approx. 0.5 miles (0.8 km) west of Toyungimpi, sample Ni 12, 5.8 ft (1.8 m) above Tapara bed; Cenomanian. (c) Pouplimanderie, sample Ni 58, 8.25 ft (2.5 m) above Tapara bed; Cenomanian. (d) Meadinga, samples Ni 98, Ni 102, and Ni 106; 12.5 ft (3.8 m), 10.5 ft (3.2 m), and 6.6 ft (2 m) above Tapara bed respectively; Cenomanian. (e) Moonkinu. Samples examined were taken from the following levels above the base of the exposed section: Ni 130, 15.8 ft (4.8 m); Ni 135, 14.2 ft (4.3 m); Ni 139, 11.2 ft (3.4 m); Ni 144, 6.6 ft (2.0 m); Ni 145, 5.9 ft (1.8 m); and Ni 147, 4.3 ft (1.3 m); Cenomanian. (f) Pulliamandera, sample Ni 192 (base of exposed section) and sample Ni 185, 14 ft (4.6 m) above sample Ni 192; Turonian. (g) Patingumputti (approx. 4 miles (6.5 km) west of Pulliamandera), sample Ni 193 (from base of exposed section) and sample Ni 198, 8.6 ft (2.6 m) above base of section; Turonian. (h) The following spot samples taken approx. 0.25 miles (0.4 km) west of Pulliamandera, and for which no strati-

graphic sequence could be established, have also been examined: Ni 122, Ni 123, and Ni 125; Cenomanian. 2. Melville Island. (a) Ant Cliff, on east coast, approx. 17.5 miles (28 km) northeast of Cape Gambier, sample Ni 280 from base of exposed section; Cenomanian. (Sample collected by Shell Development (Australia) Pty Ltd). Great Artesian Basin During the present and related studies, numerous subsurface sections have been examined from the Great Artesian Basin which incorporates the Carpentaria, Eromanga, and Surat Basins {see Day, 1964; Vine, Day, Milligan, Casey, Galloway, & Exon, 1967; Day, 1969 for stratigraphic data). The most complete mid-Cretaceous sequences available for study are of Delhi-Santos Mornington Island No. 1 well (Carpentaria Basin) and Santos Oodnadatta No. 1 well (Eromanga Basin). Mornington Island No. 1 penetrated a marine Cretaceous sequence which on data cited by Terpstra & Evans (1962) and obtained during the present study is Aptian to Cenomanian in age. Angiospermous pollen were found to be restricted to horizons (between 200 and 1,610 feet, 60.8-489.4 m) above the 'Fish Scale Zone' (Terpstra & Evans, loc. cit.) which on associated spore-pollen and dinoflagellate evidence appears to be correlative with the late middle Albian Toolebuc Limestone of the Eromanga Basin (Day, 1969). The uppermost angiosperm-bearing beds (200-1,113 ft, 60.8-338.4 m) contain Cenomanian dinoflagellate suites in which Litosphaeridium siphoniphorum and Ascodinium parvum Cookson & Eisenack, 1960 are represented. Sediments below 1,113 ft (338.4 m) are therefore of late middle Albian or late Albian age. Oodnadatta No. 1 contains a conformable sequence of marine Aptian-upper Albian strata which passes upwards into non-marine upper Albian-?Cenomanian sediments (see Freytag, 1966; Ludbrook, 1966). Angiospermous pollen have been extracted from the 87-407 ft (26.4-123.7 m) interval. The lower depth is within the Oodnadatta Formation at the top of the Coorikiana Member which may be equivalent to, or slightly younger than, the late middle Albian Toolebuc Limestone (Day,

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 1969). Horizons between 210 and 390 ft (63.8 and 118.5 m) are of late Albian age (Ludbrook, 1966). Those between 87 and 210 ft (26.4 and 63.8 m) are less securely dated although Day (1969, fig 8:6) suggested an age no younger than late Albian. A non-marine section of the Winton Formation in Haddon Downs No. 5 bore (140465 ft, 42.6-141.4 m) yielded angiospermous pollen floras. The absence of marine faunas and dinoflagellates precludes an independent means of dating the sediments, but the contained spore-pollen assemblages indicate a late Albian or early Cenomanian age (see Dettmann & Playford, 1969). Angiospermous grains are illustrated herein from the following localities in the Great Artesian Basin: 1. Queensland, Delhi-Santos Mornington Island No. 1 well; cuttings from 310-20 ft (94-7 m) and 400-10 ft (121-4 m), core 1 at 450 ft (136.8 m) and 459 ft (141.5 m). 2. South Australia, Santos Oodnadatta No. 1 well, core at 87 ft (26.4 m). 3. Haddon Downs No. 5 bore, core at 465 ft (141.4 m). Otway Basin The Otway Basin contains a thick development of sediments that spans practically the whole of the Cretaceous. The Early Cretaceous sequence is predominantly non-marine whereas marine influences are evident in much of the Late Cretaceous. Foraminiferal and palynological studies (Douglas, 1961; Dettmann, 1963; Taylor, 1964; Evans, 1966; Dettmann & Playford, 1969) have contributed to the knowledge of the depositional history of the sequence which contains several unconformities (Leslie, 1966; Reynolds, Evans, Bryan, & Hawkins, 1966). In the Otway Basin angiospermous pollen occur in late Albian and younger sediments. Late Albian occurrences are numerous and are in the largely non-marine uppermost portion of the Otway Group and its equivalents. The sediments are dated on spore-pollen evidence although a few dinoflagellates occur at some localities (e.g., F. B. H. Flaxmans No. 1 well, 7,200-20 ft; 2,189-95 m). However, species represented—Gonyaulacysta edwardsi (Cookson & Eisenack) Clarke & Verdier, 1967, Odontochitina operculata (O. Wetzel) Deflandre & Cookson, 1955—are long-ranging within the Albian and early Upper Cretaceous. Strata of probable Cenomanian age occur in the western portion of the basin in BPNL

7

Geltwood Beach No. 1 well at 3,773-93 ft (1,146-52 m). Sediments from this locality yielded a dinoflagellate suite containing Palaeohystrichophora infusorioides Deflandre, 1935, Ascodinium parvum, and probable representatives of Litosphaeridium siphoniphorum. Early Upper Cretaceous horizons examined from the eastern portion of the basin include those intersected at 7,403-8,418 ft (2,2502,558 m) in F.B.H. Port Campbell No. 2 well. On foraminiferal evidence, Taylor (1964) ascribed a Turonian age for the interval 7,4037,904 ft (2,250-2,402 m). Horizons at 7,9048,418 ft (2,402-2,558 m) are less securely dated. Dettmann & Playford (1969) suggested a late Albian-?Cenomanian to ?Turonian age for the section between 8,096-8,418 ft (2,4612,558 m). A Turonian age is here suggested for horizons at 8,096-8,188 ft (2,461-2,489 m) on the basis of the occurrence of Deflandrea acuminata. Samples that have provided figured specimens were taken from the following subsurface sections: 1. South Australia, BPNL Geltwood Beach No. 1 well, core 8, 3,771-91 ft (1,146-52 m). 2. Victoria, F.B.H. Port Campbell No. 2 well, core 7, 7,913-30 ft (2,411-7 m ) ; and core 15, 8,407-18 ft (2,555-8 m). In addition a sample from Victoria, F.B.H. Flaxmans No. 1 well, core 5, 4,479-96 ft (1,365-71 m), which contains the index and other species diagnostic of the Senonian Nothofagidites senectus Zone (of Stover & Evans, 1973), provided specimens of Australopollis obscurus (Harris) Krutzsch, 1966 that are illustrated herein by means of scanning electron micrographs. METHODS Rock samples forming the basis of this study were prepared by a method entailing immersion in 50 per cent hydrofluoric acid for 1-2 days, followed by mineral-separation process with zinc bromide solution (S.G. = 2.0). The organic residues so recovered were then subjected to brief (5-seconds) ultrasonic treatment, macerated for 2-3 minutes with Schulze solution, and finally immersed briefly in weak alkali ( < 1 per cent ammonium hydroxide). Portions of the resultant residues were mounted in unstained or lightly stained (with Safranin O) glycerine jelly under glass coverslips (No. 0) for examination by means of the light microscope. The photomicro-

Spec.Publs geoI.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

8

MARY E. DETTMANN

graphs were taken by the author on Ilford Pan F film on a Series II Zeiss Photomicroscope in the Department of Geology and Mineralogy, University of Queensland. Specimens for examination by means of the scanning electron microscope were selected by a method basically similar to that of Leffingwell, Larson, & Valencia (1970, p. 256) and Leffingwell (pers. comm.). The selected specimens were placed on a circular No. 0 coverslip (10 mm in diameter) which had previously been smeared lightly with a gelatin adhesive. After attaching the coverslip by means of an aluminium paint to a standard scanning electron microscope stub, the specimens were coated with approximately 200A of gold or gold/silver alloy in a vacuum evaporation unit. The scanning electron microscope used is a Cambridge Stereoscan IIA which was operated by Mr J. V. Hardy of the Electron Microscope Unit, University of Queensland, who also took the scanning electron photomicrographs using Kodak Tri-X film. The majority of specimens examined and photographed by means of the scanning electron microscope were retrieved and preserved as permanent mounts. The coverslips on which the specimens were initially placed were detached from the stub and inverted on to a drop of unstained glycerine jelly on glass microscope slides. Thus, these specimens became available for re-examination by means of the light microscope as the thin film of gold and gold/silver alloy coating is insufficient to mask structural and sculptural detail in transmitted light.

SYSTEMATIC SECTION In recent years considerable attention has been focussed on angiospermous pollen floras from Lower and early Upper Cretaceous strata. The pollen taxa represented in these floras are generally of small size and, as viewed under the light microscope, exhibit deceptively little complexity in exine sculpture and structure and apertural organization and configuration. Little knowledge exists as to their generic or even familial affiliations and, in consequence, most authors have treated the taxa purely as form genera, following the classification of Potonie (1960, 1966, 1970). This procedure is followed herein. Thus, generic and specific categorization of the Australian pollen types of those ages is based upon the rules of priority and typification as laid down in the International Code of Botanical Nomenclature

(Edinburgh, 1966). In following these guidelines it becomes apparent that many of the taxa proposed by van der Hammen (1956) and applicable to the Australian angiospermous pollen are of insecure validity. The situation is further confused by changes in two Articles of the Code between 1952 and 1966 (compare Arts PB3, PB6 of the 1952 Code and Arts 41, 42 of the 1961 and 1966 Codes). Potonie (1971) discussed some of the problems brought about by these changes and proposed that Article 41 of the 1966 Code be emended. Many of the generic categories about which confusion exists (mainly concerning validity) are those proposed by van der Hammen (1956) as sub-genera and subsequently raised to generic rank by Pierce (1961). Van der Hammen originally typified his sub-genera (for example, Retitricolpites, Psilatricolpites) by pollen of living species. Potonie (1960) argued that this procedure is improper and declared the taxa invalid. Pierce (1961) elevated several of the sub-generic taxa to generic rank, retaining van der Hammen's (1956) originally designated types. Van der Hammen & Wijmstra (1964), in attempting to found the genera more securely, discarded the living material as types, replacing them by fossil species. To confuse the issue still further Potonie (1966), apparently unaware of van der Hammen & Wijmstra's (1964) proposals, himself selected fossil species as types. In his 1970 publication, Potonie accepted the genera as valid and quoted what in his opinion is the correct type species for each genus. Srivastava (1969) discussed at some length the validity of one of van der Hammen's (1956) sub-genera, Retitricolpites. He argued that the original type (pollen of Neea macrophylla Poepp. & Endl.) must be retained as such, and that in consequence Retitricolpites be regarded as either a synonym or a subgeneric category of Neea. The author is in full agreement with Srivastava's opinions and further considers that other sub-generic categories based upon living material and proposed by van der Hammen (1956) are inappropriate for fossil pollen of unknown affinity. Generic and specific discrimination of the pollen types described below is based upon apertural configuration and arrangement together with sculptural and structural detail. These features are minutely expressed in many of the types, particularly in those of small size, and are difficult to assess and depict fully

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN

by means of the light microscope even at magnifications of X 1,000. Under these circumstances the scanning electron microscope has provided an excellent means for more accurately determining the surficial morphology of the pollen, and where possible descriptions are based upon observations by means of both the light and scanning electron microscopes. It has been difficult, nevertheless, to compare fully the Australian types with previously described taxa, especially those supported by brief descriptions and poor illustrations. The large number of specific epithets available (many of which have local application) for Lower and early Upper Cretaceous angiospermous pollen could well be a reflection of the limitations of the light microscope rather than real diversity of taxa (see Playford, 1971). It is anticipated that increasing use of the scanning electron microscope will enable a fuller and more objective appreciation of mid-Cretaceous angiosperm pollen morphology. The terms used in the systematic descriptions have been kept to a minimum, and are taken mainly from Erdtman (1952). Columellae (Faegri & Iversen, 1950) have been used in a wide sense to describe the sexinous rods whether they are straight-sided or have expanded tips; the precise form of the columellae is then qualified by simple terms for each species. Applications of Fischer's and Garside's Laws are in the sense of Erdtman (1952, p. 14). Thus three-aperturate grains united in a tetrahedral tetrad with the apertures meeting two and two at six points on the tetrad, have their apertures arranged according to Fischer's Law; apertures meeting three and three at four points on the tetrad are arranged according to Garside's Law. Unless otherwise specified the dimensions cited in the descriptions derive from at least 30 suitably orientated specimens. All new species are based on at least 20 adequately preserved specimens. Illustrated specimens (including those conventionally mounted after observation under the scanning electron microscope) are referred to by the preparation (prefixed D, F, K, or S)/slide number, followed by the 'N-S' and 'E-W' microscope vernier readings and registered catalogue numbers (prefixed Y) of the micropalaeontological collection of the Department of Geology and Mineralogy, University of Queensland, in which the material is deposited. The vernier readings are from Zeiss GFL Microscope (Mx 2889) of the same institution.

9

Turma PLICATES Naumova emend. Potonie, 1960 Subturma MONOCOLPATES Iversen & Troels-Smith, 1950 Genus Liliacidites Couper, 1953 Type species (by original designation): Liliacidites kaitangataensis Couper, 1953. Remarks: Judging from Couper's (1960, Plate 9, Fig. 12) illustration of the type species and from an examination of Australian grains closely comparable to the type species, Liliacidites is characterized by a differentially thickened exine. The thickened areas of exine occur at and about the equator in areas circumscribing the ends of the short equatorial axis of the grain. The presence of a differentially thickened exine distinguishes the genus from Clavatipollenites Couper, 1958 (see also discussion under generic heading of Clavatipollenites) . A further feature which is used herein for discrimination of the two genera is the form of the sulcus; in Liliacidites the sulcus appears to be developed in both the nexine and sexine, whilst in Clavatipollenites the sexine may be continuous over the nexine sulcus. Couper (1953, 1960) described species of Liliacidites as having a surface reticulum of variable mesh-size, the smaller meshes occurring at the ends of the grain and along sulcal margins {i.e. in areas of thin exine). In Australian examples, the surface reticulum is of ± uniform mesh-size, although the luminal areas below the surface do exhibit a variation in size as described by Couper. Affinity: Couper (1953) suggested a liliaceous affinity. Monosulcate grains of the Liliaceae usually are derived from tetragonal tetrads with the aperture on the distal surface (Erdtman, 1952, 1969), features which have not been determined for the Australian Cretaceous representatives of Liliacidites. It is of interest to note that some of the grains allocated herein to Liliacidites show a tendency to rupture on the surface diametrically opposite the sulcus; superficially this rupture simulates a second sulcus. Other extant plants that produce pollen similar to Liliacidites occur in the Amaryllidaceae (see Erdtman, 1952, 1969). Liliacidites cf. kaitangataensis Couper, 1953 (Plate 1, Figs 1-5) Description: Pollen grains free, anisopolar, monosulcate, bilateral, ellipsoidal; amb elliptical. Sulcus long, developed in both nexine and

Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

MARY E. DETTMANN 10 sexine, extending to equator, parallel to long assuming the form and dimensions of the ?disaxis of grain, often ragged and with rounded tal sulcus (Plate 1, Fig. 1). equatorial extremities. Exine two-layered, conDistribution: Of persistent occurrence in sisting of a thin nexine (ca 0.2/* thick) and a the Cenomanian of Bathurst and Melville thicker differentially thickened, columellate Islands. Couper (1953, 1960) reported L. sexine. Sexine thickest (2-2.5/* thick) at equa- kaitangataensis from New Zealand upper tor in areas circumscribing ends of short axis Senonian and Maestrichtian with doubtful of grain, thinnest (0.75-1/*) in a continuous occurrences in the Eocene. longitudinal band that includes sulcal margins and equivalent areas on diametrically opposite Liliacidites cf. intermedins Couper, 1953 surface. Columellae with irregular bases and ex(Plate 1, Figs 6-8) panded heads; largest (2/* high, 1/* basal diaDescription: Pollen grains free, anisopolar, meter, 1.5-2/* head diameter) and more widely- monosulcate, ellipsoidal; amb ellipspaced (0.75-1.5/*) in regions where exine is tical. Sulcus bilateral, developed in both sexine and thickest; becoming smaller and more closely nexine, orientated parallel to long axis of grain, spaced in areas of thin sexine. Columellate irregular in outline, extending to heads fused to form a surface reticulum with equator. Exine 1-1.5/* thick, with aalmost thin (ca muri 1.5-2/* wide and subcircular lumina 1-2/* /* thick) nexine and a thicker, differentially diameter. In well-preserved specimens surface 0.1 columellate sexine. Sexine thickest reticulum of uniform mesh-size in all regions thickened, at equator in areas circumscribing ends of grain (in contrast to marked differentiation of(1.5/*) short axis of grain; thinnest (0.75-1/*) in a in size and spacing of supporting columellae). continuous longitudinal band that includes In corroded specimens muri of reticulum may sulcal margins and equivalent area on diametribe partially to completely broken down and opposite surface. Columellae with cylinthus lumina approximate dimensions of spaces cally drical columns and expanded heads; between columellae (i.e. lumina are largest in largest (1/* high, 0.4/* basalflat-topped 0.6/* areas of thick sexine and smallest in regions head diameter) and more widelydiameter, spaced (0.5of thin sexine). Muri of reticulum mostly uni- 1/*) in areas of thick sexine; smaller and more columellate, occasionally duplicolumellate. closely-spaced in areas of thin sexine. ColuDimensions: Equatorial diameter; length 38 mellate heads fused to form surface reticulum (52) 65/*, breadth 20 (34) 50/*. Polar dia- with muri 0.5/* wide and circular to polygonal lumina 0.5-1/* diameter. In well-preserved meter 25 (36) 48/x. lumina ± uniform in size in all Remarks and comparison: The grains have specimensof grain; in corroded examples lumina a surface reticulum of ± uniform mesh-size, regions sometimes largest in areas of thick sexine. Muri a feature detected by means of the scanning electron microscope (Plate 1, Fig. 5). Couper of reticulum unicolumellate. Dimensions: Equatorial diameter; length 27 (1953, p. 56) described his species Liliacidites kaitangataensis with lumina that 'vary in dia- (36) 53/*, breadth 17 (26) 36/*. Polar diameter from 5/* at centre of grain to 1/* at ends meter 20 (25) 35/*. of grain'; it seems possible that these dimenRemarks and comparison: The grains appear sions represent the spacing of the columellae to be similar to Liliacidites intermedins Couper, beneath the surface reticulum. A similar varia- 1953 but have a thinner exine and are generally tion in mesh-size has been observed in corroded smaller. Couper (1953, p. 56) described the examples (in which the surface reticulum is surface reticulum as having lumina of variapartially broken down) recovered from Aus- ble size (2-3/* near centre of grain, ca 1/* at tralian deposits. However, the Australian grains ends of grain) although it is possible that are only compared with Couper's species as these dimensions represent the spacing of the they have a thicker exine and display a greater columellae beneath the surface reticulum. variation in size-range. In organization L. cf. intermedins is similar The exine of grains here attributed to L. cf. to L. cf. kaitangataensis, but is distinguishable kaitangataensis is clearly of differential thick- in its thinner exine, more slender and more ness. The thin areas of exine occur about the closely-spaced columellae, and smaller size. sulcal margins and in an equivalent area on As in L. cf. kaitangataensis the exine is of the diametrically opposite (?proximal) face; differential thickness and a rupture assuming in corroded examples this thin area of ?proxi- the form of the ?distal sulcus may be demal exine is sometimes ruptured, the rupture veloped on the ?proximal surface.

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN The grain 'monosuicate type 1' illustrated by Doyle (1969, Plate 5, Figs 1, m) resembles L. cf. intermedins. Distribution: Found persistently in the Cenomanian and infrequently in the Turonian of Bathurst and Melville Islands. L. cf. intermedins is also represented in the Cenomanian of Mornington Island. Couper (1953, 1960) reported L. intermedins as ranging from Maestrichtian to Middle Miocene in New Zealand. Genus Clavatipollenites

11

Affinity: The possible angiospermous nature of Clavatipollenites has been discussed and debated by many authors (see Couper, 1958; Brenner, 1963; Kemp, 1968; Doyle, 1969; Muller, 1970). Muller and Doyle accepted an angiospermous affinity for the genus and implied that the aperture is distal, a character that can be confirmed only from material united in tetrads. The possible derivation of isopolar, in particular tricolpate, forms from Clavatipollenites by way of Asteropollis has also been suggested (Doyle, 1969; Muller, 1970; and cf. Chaloner, 1970; this paper).

Couper, 1958

Type species (by original designation): Clavatipollenites hnghesii Couper, 1958.

Clavatipollenites sp. (Plate 2, Figs 8-10)

Remarks: Kemp (1968) clarified the morphology of Clavatipollenites Couper, 1958 and concluded that the genus and Liliacidites are morphologically similar. She suggested that the two genera be retained; Clavatipollenites for mid-Cretaceous forms of uncertain affinity and Liliacidites for Upper Cretaceous and Tertiary grains of angiospermous (possibly liliaceous) origin. As noted above two distinct categories of reticulate, monosuicate grains have been recognized in the Australian material. One type, here referred to Clavatipollenites, is characterized by a uniformly thick exine and a sulcus that is invariably developed in the nexine, but only occasionally developed in the sexine, as a result of a breakdown of the columellate processes in an irregular area broadly corresponding to the nexine sulcus. The second type, allocated above to Liliacidites, has an exine of differential thickness and a sulcus that is clearly and regularly formed in both nexine and sexine. Thus, distinction between Liliacidites and Clavatipollenites can be achieved on a morphological basis.

Description: Grains free, anisopolar, monosuicate, subspheroidal; amb subcircular. Exine two-layered, consisting of nexine, 1-1.5^ thick, and an outer, columellate sexine, thick. Columellae 0.75-1^ high, with slender cylindrical columns and expanded tips; free or fused in apertural region to form an imperfect reticulum which may be partially or wholly broken-down to expose sulcus. In other regions of grain, columellae fused to form a regular reticulum with polygonal lumina 0.5^ diameter. Sulcus developed in nexine as an elongated slit that is broadest in central regions, tapering towards extremities. Sexine entire over sulcus or with an irregular rupture formed from a breakdown of columellate processes.

Clavatipollenites resembles closely Asteropollis Hedlund & Norris, 1968 and there seems little doubt that the two genera have a closely related origin. In both the aperture is clearly and regularly expressed in the nexine but the sexine may be entire or irregularly ruptured in the sulcal area. Since the irregular sexinous rupture of Asteropollis may assume a shape distinct from the nexine sulcus, discrimination between it and Clavatipollenites must be based upon the shape of the nexine sulcus. In Clavatipollenites the nexine aperture is monosuicate, whilst in Asteropollis a three- to fivebranched aperture is represented.

Dimensions: Grain length 17 (20) 26^ breadth 15 (19) 25^; depth (3 specimens) 11 (17) 19/x. Remarks and comparison: The Australian specimens are similar in sculptural and sulcal features to the type species, Clavatipollenites hughesii Couper, 1958, with which they may be conspecific. However, the Australian specimens are more robust and are consistently subcircular in equatorial outline. C. rotundns Kemp, 1968 is distinct in having a darkened zone (?thickening) adjacent to the sulcus; C. minntus Brenner, 1963 differs in its smaller size. Distribution: Of rare occurrence in the eastern Australian Cretaceous. It is known from latest Albian and Cenomanian of the Great Artesian Basin; Cenomanian-Turonian of Bathurst and Melville Islands, and probable Cenomanian-Turonian of the Otway Basin. Its incoming in all sections examined postdates the appearance of tricolpate angiospermous grains.

Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

12

MARY E. D

Subturma TRIPTYCHA Naumova, 1939 Genus Cupuliferoidaepollenites Potonie, Thomson, & Thiergart, 1950 Type species (by original designation): Cupuliferoidaepollenites liblarensis Thomson in Potonie, Thomson, & Thiergart, 1950. Remarks: Tricolpate, isopolar pollen grains having a smooth or faintly scabrate exine have been variously assigned to Cupuliferoidaepollenites Potonie, Thomson, & Thiergart, 1950, Tricolpopollenites Thomson & Pflug, 1953, Cornaceoipollenites Potonie, 1951, and Psilatricolpites van der Hammen, 1956. Tricolpopollenites and Cornaceoipollenites are now known to be based upon tricolporate grains (Krutzsch, 1959, p. 43) and Psilatricolpites has, as its type, pollen of the extant Bartsia santalinaefolia (H. B. K.) Benth. and is considered here to be a junior synonym of Bartsia. Potonie (1966) attempted to conserve Psilatricolpites for fossil tricolpate grains by selecting Pierce's (1961) species P. psilatus as the type. Arguments against the validity of this procedure have been documented by Srivastava (1969, p. 56) in the parallel case of Retitricolpites van der Hammen, 1956 (see also introductory remarks to Systematic Section of this paper). Cupuliferoidaepollenites cf. parvulus (Groot & Penny) comb, no v. (Plate 2, Figs 11-15) 1960 Tricolpopollenites parvulus Groot & Penny, p. 232, Plate 2, Figures 8, 9. 1967 Psilatricolpites parvulus (Groot & Penny) Norris, p. 107, Plate 17, Figures 5-7. Description: Pollen grains free or united in tetrahedral tetrads, isopolar, tricolpate, prolate to subspheroidal; amb subcircular to convexly subtriangular. Colpi straight-sided narrow slits, extending almost to poles; arranged according to Fischer's Law as observed in examples united in tetrads. Exine 1-1.5^ thick, with a smooth or faintly scabrate surface; sexine and nexine not clearly differentiated. Sexine without visible columellae at magnifications of X 1000. Dimensions: Equatorial diameter 7 (10) 13/x; polar diameter 10 (12) 14/x. Remarks and comparison: The Australian pollen grains are similar to Cupuliferoidaepollenites parvulus (Groot & Penny) comb. nov. but have longer colpi. Tricolpites erugatus Hedlund, 1966 has long colpi but differs from

the Australian pollen of C. cf. parvulus in displaying better defined scabrate sculpture and a thinner exine. Grains figured by Doyle (1969, Plate 3, Figs a, b) may well be comparable to C. cf. parvulus. Distribution: Recovered from Cenomanian and Turonian of northern Australia (Bathurst, Melville, and Mornington Islands). The similar grains of C. parvulus have been reported widely from late Albian-Turonian of North America (Groot & Penny, 1960; Groot, Penny, & Groot, 1961; Brenner, 1967; Norris, 1967; Singh, 1971). Genus Tricolpites Cookson ex Couper, 1953 Type species (by subsequent designation of Couper, 1953, p. 61): Tricolpites reticulatus Cookson, 1947. Remarks: The genus is used here in the sense of Potonie (1960, p. 95) and Srivastava (1969, p. 55) for the reception of tricolpate, isopolar grains having simple slit-like colpi and a surface reticulum of ± uniform development and of mesh-size 1-2^ or less. These characters are exhibited by the type species Tricolpites reticulatus Cookson, 1947, which has been re-examined under both the light and scanning electron microscopes (Cookson, Dettmann, & Playford, in prep.). Other generic categories designed to incorporate finely and uniformly reticulate, tricolpate species have been found to be invalid (Retitricolpites van der Hammen, 1956; see Kemp, 1968, p. 430; Srivastava, 1969, p. 55; and cf. Potonie, 1966, p. 163; 1970, p. 110), inappropriate (Tricolpopollenites Thomson & Pflug, 1953; see Potonie, 1966, p. 168), or of uncertain morphology (Foveotricolpites Pierce, 1961). Two of the species recorded herein, Tricolpites minutus (Brenner) comb. nov. and T. cooksonae sp. nov., are represented by occasional tetrahedral tetrads, with the apertures arranged according to Fischer's Law. Tricolpites minutus (Brenner) comb. nov. (Plate 4, Figs 1-4) 1963 Tricolpopollenites minutus Brenner, p. 93, Plate 40, Figures 5, 6. 1967 Cornaceoipollenites minutus (Brenner) Norris, p. 107, Plate 17, Figures 7-11. 1971 Cupuliferoidaepollenites minutus (Brenner) Singh, p. 194; Plate 29, Figures 8, 9. 1971 Tricolpites micromunus (Groot & Penny) Burger, p. 8, Plate 1, Figures 5, 7.

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 13 Description: Pollen grains free or occasion- pate, subspheroidal to oblate, free or occaally united in tetrahedral tetrads, isopolar, tri- sionally united in tetrahedral tetrads; amb subcolpate, prolate to sub-prolate; amb subcir- circular. Colpi narrow parallel-sided slits cular to convexly subtriangular (compressed formed in both sexine and nexine; arranged specimens). Colpi long (at least 3/4 polar according to Fischer's Law; length %-% polar diameter of grain), narrow, ± parallel-sided diameter of grain. Exine two-layered; nexine slits developed in both nexine and sexine; 0.75-1/* thick, without discernible structure arranged according to Fischer's Law. Exine under light microscope at magnifications of X 0.75-1/* thick, clearly differentiated into nexine 1,000. Sexine as thick as nexine, columellate. and sexine which are approximately equal in Columellae 0.75/t high, spaced 0.2-0.6/* apart, thickness. Nexine appears homogeneous at with slender cylindrical columns (ca 0.1 /* basal magnifications of X 1,000; sexine distinctly diameter) and expanded subglobular heads columellate and with a microreticulate sur- (0.3/* diameter). Columellate heads fused at face pattern. Columellae slender, closely- their bases in groups of 12-15 to form muri spaced, fused at their tips to enclose circular 0.4/* wide which enclose subcircular to polylumina ca 0.2-0.4/x in diameter; reticulum of gonal lumina; small and large lumina (ranging uniform development in all regions of grain. from 0.4-1.8/* diameter) randomly distributed Dimensions: Equatorial diameter 7 (12) over surface of grain. Crests of muri appear granular at high magnifications (X 3,000), 15fx; polar diameter 10 (13) 16/x. Remarks and comparison: The Australian the granules representing the distal extremities grains conform with Brenner's (1963) diag- of the columellate heads above the surface of nosis and appear identical with the specimen the muri (see Plate 4, Figs 17-19). figured by Doyle (1969; Plate 2, Figs c, d) Dimensions: Equatorial diameter 14 (18) which clearly shows the surface reticulum. polar diameter (3 specimens) 15 (17) This character suggests that the species may be 20/*; 18/*. more appropriately placed in Tricolpites rather Holotype: Preparation S340/2, 16.1 90.5, than Cupuliferoidaepollenites Potonie, Thomson, & Thiergart, 1950. Specimens described Y. 1472; Plate 4, Figures 13-15. Polar aspect. and illustrated by Burger (1971, p. 8; Plate 1, Amb subcircular, 17/* diameter. Colpi 11/* long, Figs 5, 7) as Tricolpites micromunus (Groot gaping at equator, margins smooth. Exine 1.8/* & Penny) Burger, 1971 are almost certainly re- thick; sexine 0.9/* thick, columellate and with presentatives of Tricolpites minutus (Brenner) a reticulate surface. Slender columellae 0.75/* comb, no v.; they are distinct from T. micro- high, bases ca 0.1/* diameter, expanded heads munus which has coarser and more angular 0.3/* diameter; heads fused to form surface reticulum with polygonal lumina 0.5-1.7/* dialumina. The species bears a superficial resemblance to meter. Australian grains allocated to CupuliferoidaeType locality: Bathurst Island, Meadinga; pollenites cf. parvulus (Groot & Penny) comb, sample Ni 102. Cenomanian. nov., but are distinct in having a thinner Remarks and comparison: Almost all exexine which is finely reticulate. T. albiensis amples recorded glycerine jelly mounts are Kemp, 1968 is similar to T. minutus, differing orientated in polarin aspect. Lateral aspects have only in having a thicker exine (as illustrated been viewed under the scanning electron microby Kemp, 1968, Plate 81, Figs 1-22). scope and illustrations on Plate 4, Figures 17, Distribution: T. minutus appears to be con- 18 depict a specimen in both polar and lateral fined to the Cenomanian on Bathurst and Mel- views. ville Islands and is known from late AlbianTricolpites cooksonae sp. nov. is similar to early Cenomanian of the Great Artesian Basin (present study; Burger, 1971). Extra-Aus- grains assigned by Singh (1971) to Retitricoltralian records include Albian and Ceno- pites virgeus (Groot, Penny, & Groot) Brenner, manian of North America (Brenner, 1963; 1963. The holotype of the latter species is distinct, however, in its larger size, longer colpi, Norris, 1967; Doyle, 1969; Singh, 1971). and more robust muri. Retitricolpites vulgaris Tricolpites cooksonae* sp. nov. Pierce, 1961 and Tricolpites sp. 2 of Kemp (Plate 4, Figs 13-19) (1968) have longer colpi, thinner exine, and Description: Pollen grains isopolar, tricol- smaller lumina; Tricolpopollenites platyreticu* The species is named for Dr I. C. Cookson. Spec.Publs geol.Soc.Aust, 4: pp. 3-34, Pis 1-6, 1973.

MARY E. DETTMANN 14 latus Groot, Penny, & Groot, 1961 is described Basin, and in Cenomanian-Turonian of the Otway Basin and Bathurst Island. as having larger lumina. Distribution: Of infrequent occurrence in Genus Rousea Srivastava, 1969 Cenomanian and Turonian of Bathurst and Melville Islands. The similar grains described Type species (by original designation): by Singh (1971) as Retitricolpites virgeus are Rousea subtilis Srivastava, 1969. from middle Albian-Cenomanian of western Remarks: The genus is based upon the Canada. Late Cretaceous species, Rousea subtilis Srivastava, 1969, and is characterized by tricolpate Tricolpites sp. pollen having a surface reticulum, the mesh(Plate 4, Figs 5-12) size of which is largest in the mesocolpia and Description: Pollen grains isopolar, tricol- smallest in the apocolpia. These features are pate, prolate to subspheroidal; amb subcircular. exhibited by the mid-Cretaceous pollen here Colpi at least 3/4 polar diameter, developed attributed to Rousea. Other mid-Cretaceous in both nexine and sexine as narrow, parallel- forms that appear to conform with Rousea insided slits. Exine 1-1.5/* thick; nexine 0.5/x clude Retitricolpites prosimilis Norris, 1967 thick, without discernible structure at magni- and pollen type 3 of Doyle (1969; Plate 2, fications of X 1000. Sexine thicker (1 /*) than Figs n, o). nexine, distinctly columellate and with a surAffinity: Srivastava (1969) compared his face reticulum. Columellae 0.75/x high, spaced Late Cretaceous R. subtilis with pollen of the 0.75-1/* apart, with cylindrical columns (0.2/* Salicaceae, familial alliance may not basal diameter) and expanded subglobular be applicablebutto this heads (0.5/* diameter). Columellate heads con- scribed below. the mid-Cretaceous pollen denected at their bases by low bridge-like muri 0.25-0.4/* wide. The columellate heads, which Rousea georgensis (Brenner) comb. nov. project above the bridge-like muri, are con(Plate 2, Figs 16-17) nected in groups of 4-7 to enclose polygonal lumina; small and large lumina (ranging from 1963 Retitricolpites georgensis Brenner, p. 91; 0.2 to 1.25/JL diameter) randomly distributed Plate 38, Figure 6. over entire surface. 1971 Tricolpites georgensis (Brenner) Burger, Dimensions (15 specimens): Equatorial p. 7; Plate 3, Figures 2, 4. diameter 12 (16) 22/*; polar diameter 16 (18) Description: Pollen grains tricolpate, iso25//,. polar, prolate to subprolate; amb circular. Remarks and comparison: Scanning electron Colpi long (at least 3/4 polar diameter), paralmicrographs (Plate 4, Figs 9-12) illustrate the lel-sided slits which are developed in both surface features of the grains described above. nexine and sexine. Exine 1.5/* thick; nexine The columellate heads are fused at their bases, 0.5/* thick, thinner than sexine (1/*) which is and the distal extremities of the heads project distinctly columellate. Columellae 0.75/* high, above the connecting muri. with cylindrical columns and expanded subTricolpites sp. resembles Foveotricolpites globular heads that are fused to form a surface concinnus Singh, but judging from the illus- reticulum. Reticulum composed of muri 0.75/* trations (1971; Plate 29, Figs 10, 11) the Aus- wide and polygonal lumina that are largest tralian grains are smaller and have a finer- (0.75 — 1.5/* diameter) in the mesocolpal meshed surface reticulum. Other species that areas, decreasing to ca 0.1/* at poles and along may be similar to Tricolpites sp. include Reti- colpal margins. tricolpites sphaeroides Pierce, 1961 and R. Dimensions (10 specimens): Equatorial diaoblatoides Pierce, 1961; detailed comparisons meter 11 (15) 18/*; polar diameter 15 (20) must await a clearer morphological under- 24/*. standing of Pierce's species. Comparison: The grains described here are Tricolpites sp. is distinct from T. cooksonae smaller, but in other respects are comparable which has more slender columellae, shorter to Rousea georgensis as described and illuscolpi, and a thicker exine; T. minutus has a trated by Brenner (1963). The species is disconsiderably finer-meshed surface reticulum. tinct from R. subtilis Srivastava, 1969 which is Distribution: Of infrequent occurrence in larger, with a thicker exine and a coarserlate Albian-Cenomanian of the Great Artesian meshed reticulum. Retitricolpites prosimilis

ANGIOSPERMOUS POLLEN ] ROM ALBIAN TO TURONIAN Norris, 1967 has a finer-meshed reticulum and pollen type 3 of Doyle (1969; Plate 2, Figs n, o) has thickenings along the margins of the colpi. Retitricolpites fragosus Hedlund & Norris, 1968 exhibits differential development of the surface reticulum, but the lumina are largest in the polar regions. Distribution: R. georgensis is of widespread distribution in the Albian and Cenomanian of North America (Brenner, 1963; Norris, 1967; Hedlund & Norris, 1968; Singh, 1971; Playford, 1971). The species occurs infrequently in middle-upper Albian and Cenomanian of the Great Artesian Basin, in the Cenomanian of Bathurst and Melville Islands, and in late Albian and Cenomanian of the Otway Basin (Burger, 1971; present study). Genus Striatopollis Krutzsch, 1959 Type species (by original designation): Striatopollis sarstedtensis Krutzsch, 1959. Remarks: Several genera have been proposed for tricolpate pollen having a finely striate exine. Striatopollis Krutzsch, 1959 is based upon the Tertiary species S. sarstedtensis Krutzsch, 1959 and is the senior synonym of Striopollenites Rouse, 1962 (see Potonie, 1966, p. 164). Similar grains have also been assigned to Striatricolpites van der Hammen, 1956 which is invalid (Potonie, 1970). Although the majority of species described under these generic categories are of Early Tertiary age (see Krutzsch, 1959; Rouse, 1962; Germeraad, Hopping & Muller, 1968), it is now evident that morphologically similar pollen have wide distribution in mid-Cretaceous sediments (Groot & Groot, 1962; Jardine & Magloire, 1965; Norris, 1967; Hedlund & Norris, 1968; Singh, 1971; present study). Affinity: Morphologically similar pollen are known in the Fabaceae, for example in Crudia amazonica Spruce (see Germeraad, Hopping, & Muller, 1968, Plate 12, Figs 5, 6). Striatopollis cf. paraneus (Norris) Singh, 1971 (Plate 2, Figs 18-22) 1967 Retitricolpites paraneus Norris, p. 109; Plate 18, Figures 15-20. 1971 Striatopollis paraneus (Norris) Singh, p. 206; Plate 32, Figures 1-3. Description: Pollen grains isopolar, tricolpate, prolate; amb subcircular to circular. Exine In thick, two-layered, nexine thin (ca 0.2/x thick); sexine thicker (0.75^), with a sur-

15

face pattern of ± parallel, occasionally anastomosing and bifurcating muri which show a preferred orientation parallel to the polar axis of the grain. Muri 0.4-0.5^ wide and high, with secondary cross striations (ridges) that are evenly spaced 0.1-0.2^ apart. Exinal ornament thus has a 'ropy' appearance visible only at magnifications of at least x 3,000; at lower magnifications, the secondary cross striations are not visible but the primary muri appear blurred and seemingly cannot be focused sharply. Sexine in colpal regions with two to three rows of free standing columellae which may interlock over colpi; columellae 0.3-0.4^ high, with expanded subcircular heads 0.4-0.5/x diameter. Colpi developed in nexine as narrow, parallel-sided slits, 2 / 3 length of polar axis. Dimensions: Equatorial diameter 10 (12) 1 5/JL; polar diameter 13 (15) 16^. Remarks and comparison: The fine sculptural details of the grains here compared with Striatopollis paraneus (Norris) Singh, 1971 were determined from observations under the scanning electron microscope. The sexine wall structure has not been determined precisely although present data suggest that the sexine is columellate, the fused columellae forming the rope-like surface ridges. As observed under the light microscope, the Australian grains exhibit a remarkable similarity to, and may be conspecific with, S. paraneus as illustrated by Norris (1967) and Singh (1971). Conspecificity can only be established when the precise form of the muri and the sculptural detail in the colpal regions of the North American specimens have been determined. Grains allocated to S. paraneus by Hedlund & Norris (1968) are distinct in having a thicker exine and coarser muri. Retitricolpites vermimurus Brenner, 1963, Striopollenites dubius Jardine & Magloire, 1965, and Groot & Groot's (1962; Plate 9, Figs 11-14) specimens of Striatopollis sarstedtensis Krutzsch, 1959 have more robust muri, wider colpi, and thicker exine. Striatricolpites catatumbus Gonzalez, 1967 is larger and has longer colpi and a thicker exine. Muller's (1968) species Striatricolporites conspicuus and S. minor are described as tricolporate. Distribution: Of rare occurrence in the Cenomanian of Bathurst and Mornington Islands. The similar form Striatopollis paraneus has been reported from middle-upper Albian of western Canada (Norris, 1967; Singh, 1971; Playford, 1971).

Spec.Publs geol.Soc.Aust, 4: pp. 3-34, Pis 1-6, 1973.

16

MARY E. DETTMANN

Genus Phimopollenites* gen. nov. Type species (here chosen): Phimopollenites pannosus (Dettmann & Playford) comb. nov. Diagnosis: Pollen grains tricolpoidate, isopolar, oblate to prolate. Exine differentiated into nexine and sexine, sexine columellate. In mesocolpal and polar areas columellae fused at their distal extremities to form foveolate to finely reticulate surface pattern. In colpal areas columellae free-standing, interlocked over nexine colpoids to form elliptical 'operculoid' membranes. Operculoid membranes may be partially to completely broken down to expose broadly elliptical nexine colpoids. Surface reticulum of uniform development with meshes up to 2/x diameter. Comparison: The genus is similar to Tricolpites Cookson ex Couper, 1953 in its uniformly developed surface reticulum but differs in its colpal characters. In Tricolpites, the colpi have smooth margins and are expressed as ± parallel-sided slits in both nexine and sexine; in Phimopollenites gen. nov., the sexine aperture is elliptical and is formed as a result of a breakdown of an operculoid membrane which consists of free-standing columellae. Dactylopollis Muller, 1968 is distinct in having a striate sexine in polar regions and in lacking operculoid membranes in the colpal regions. Affinity: Many extant species produce pollen morphologically similar to Phimopollenites. In particular pollen of Fothergilla Murr. (Hamamelidaceae) and Tetracentron Oliver (Tetracentraceae) resemble fossil grains here attributed to Phimopollenites pannosus (Dettmann & Playford) comb. nov. and P. augathallaensis (Burger) comb. nov. Phimopollenites pannosus (Dettmann & Playford) comb. nov. (Plate 3, Figs 1-11) 1968 Tricolpites pannosus Dettmann & Playford, p. 84; Plate 8, Figures 5-8. 1971 Tricolpites variabilis Burger, p. 8; Plate 1, Figures 1, 4. 1971 Psilatricolpites pannosus (Dettmann & Playford) Burger, p. 6 (?grains illustrated on Plate 1, Figures 2, 3, 6.) Description: Pollen grains isopolar, tricolpoidate, free or occasionally united in tetrahedral tetrads, prolate to subspheroidal; amb * Gk., phimos, muzzle.

trilobed, fossaperturate, subcircular in flattened specimens. Exine two-layered; composed of an inner nexine 0.15^ thick and a columellate sexine 0.5-0.75/x thick. Columellae ca 0.4^ high, with expanded heads ca 0.3jx diameter, barely visible in optical sections at x 1,000 magnifications; fused to form a microreticulate surface pattern except in colpal regions where they are free-standing. Reticulum ± uniformly developed, with muri 0.2-0.3/x wide and polygonal to elongated lumina up to 0.6/x diameter. Colpoids polar diameter of grain, developed in nexine in concave regions of grain, elliptical in outline, 3-4^ wide at equator, tapering towards polar extremities which are rounded; often partially to completely covered by an operculoid sexinous membrane which corresponds in size and position to nexine colpoid. Operculoid membrane with free-standing columellae 0.3-0.4^ high, 0.20.3/x head diameter; membrane may be partially to completely broken down to reveal nexine colpoid. Apertures arranged according to Fischer's Law as seen in tetrahedral tetrads. Dimensions: Equatorial diameter 9 (17) 25/x; polar diameter 13 (22) 28/x. Remarks and comparison: The species was previously regarded as faintly scabrate to psilate (Dettmann & Playford, 1968), but with more sophisticated optical equipment (Nomarski interference contrast, scanning electron microscope) the surface pattern is now known to be microreticulate. The apertures are covered by finely granulate operculoid membranes that may be irregularly ruptured to expose colpoids. Reexamination of the holotype and other specimens figured by Dettmann & Playford (1968, Plate 8, Figs 5-8; and cf. this paper, Plate 3, Figs 1-4) illustrates that Phimopollenites pannosus (Dettmann & Playford) is comparable to Burger's (1971, p. 84; Plate 1, Figs 1, 4) Tricolpites variabilis. The specimens Burger (loc. cit., p. 6; Plate 1, Figs 2, 3, 6) referred to P. pannosus are possibly correctly identified although the surface reticulum appears less conspicuous. P. pannosus resembles Tricolpites albiensis Kemp, 1968 but is larger and has shorter colpi which have operculoid membranes. Distribution: A widely distributed species in eastern Australia, where it is one of the earliest appearing 'tricolpate' species with initial occurrences in late middle Albian of the Great Artesian Basin (Burger, 1971; this study). The

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN

species is sometimes abundant in the Winton Formation (late Albian-?Cenomanian) of the Great Artesian Basin and in its initial occurrences in probable late Albian sediments of the Otway Basin. In the latter area it ranges into probable Turonian strata. At Bathurst and Melville Islands P. pannosus occurs infrequently in the Cenomanian but has not been observed in Turonian sequences.

uniform in surface sculptural features. Although the species is typically tricolpoidate, several examples (Plate 2, Fig. 16) recovered are tetracolpoidate. Phimopollenites augathallaensis (Burger) comb. nov. is distinct from P. pannosus in its larger size and more coarsely sculptured sexine. Tricolpites sagax Norris, 1967 shows some resemblance but appears to lack the operculoid membranes of free standing columellae in the Phimopollenites augathallaensis (Burger) comb, colpal areas. nov. Distribution: Burger (1971) described the species from the middle-upper Albian and (Plate 3, Figs 12-17) 1971 Tricolpites augathallaensis Burger, p. 7; ?Cenomanian of the Eromanga Basin. In the present study P. augathallaensis has been rePlate 2, Figures 1, 2. covered in small numbers from the CenoDescription: Pollen grains isopolar, tricol- manian Bathurst Island and more frequently poidate, prolate to subspherical; amb subcir- from theof Cenomanian of Mornington Island cular. Exine two-layered; composed of an inner and upper Albian-?Cenomanian of South Ausnexine 0.5^ thick and a distinctly columellate tralia. sexine 1.5/* thick. Columellae 0.5-0.75^ high, with cylindrical columns 0.3/x diameter and exSubturma POLYPTYCHES Naumova, 1939 panded subglobular heads 0.4-0.6^ diameter; Genus Asteropollis Hedlund & Norris, 1968 heads fused to form a fine surface reticulum. Type species (by original designation) : Muri with rounded crests 0.5-0.75/x wide, enclose subcircular lumina 0.2-0.4^ diameter; Asteropollis asteroides Hedlund & Norris, reticulum uniformly developed in mesocolpal 1968. and polar areas. Colpoids formed in nexine Remarks: As discussed under the generic as elliptical openings which are widest at of Clavatipollenites, Asteropollis Hedequator, tapering towards pointed polar ex- heading lund & Norris, 1968 and Clavatipollenites are tremities; length 2/3 polar diameter. Nexine morphologically similar, differing only in the colpoids covered with operculoid membranes shape of the nexine sulcus. The genus is monocomposed of free-standing sexinous columellae typic, although is seems certain that grains rewhich may be interlocked in the form of a ferred by Singh (1971) Liliacidites trichozip-fastener. Columellae distinctly visible under tomosulcatus and some ofto those described by light microscope at magnifications of x 1,000; Groot & Groot (1962) as Apiculatisporis 1 p high and with subglobular heads 0 . 2 - i n garis are representatives of distinct speciesvulof diameter; often partially broken down to re- Asteropollis. veal nexine colpoids. Following Hedlund & Norris (1968) the Dimensions: Equatorial diameter 22 (23) genus is here referred to Subturma Polyptyches Naumova, 1939, although the representation 45^; polar diameter 22 (35) 40^. of a three-branched sulcus in types referable to Remarks and comparison: A distinctive type Asteropollis suggests an equally appropriate that incorporates the largest 'tricolpate' angio- inclusion within Subturma Trichotomonospermous grains extracted during the present investigation. The specimens described here sulcates Erdtman, 1945. broaden the size range as quoted in the diagnosis (Burger, 1971, p. 7). Burger described Asteropollis asteroides Hedlund & Norris, 1968 the surface reticulum with lumina larger (Plate 2, Figs 1-7) (0.3-0.8/x) than the enclosing muri (0.2-0.3/x), Description: Grains anisopolar, oblate to but these dimensions probably represent the subspheroidal with a circular amb; tri- to dimensions of the columellae and the inter- pentachotomosulcate ?distally. Sulcus with vening spaces at the base of the columellate three to five subequal ± parallel branches layer rather than at the outer surface of the (each 2-4^ wide) which to equator sexine. The five specimens (taken from diff- where they broaden to formextend rounded extremierent localities) examined by means of the ties. Exine 2-2.5//, thick; inner unsculptured scanning electron microscope are remarkably nexine 1-1.5^ thick, thins considerably at aperSpec.Publs geal.Soc.Aust, 4: pp. 3-34, Pis 1-6, 1973.

18

MARY E. I

tural margins and is absent in apertural regions. Sexine 1-1.5p thick, columellate; columellae lju, high, with slender cylindrical columns (0.2^ in diameter) and expanded tips (0.4/x in diameter). In apertural regions columellae discrete or fused in irregular clumps to form an imperfect reticulum which may be partially or wholly broken-down to expose nexine sulcus. In non-apertural regions, columellae fused to form a regular reticulum with subcircular to polygonal lumina 0.5^ diameter. Dimensions'. Equatorial diameter 15 (21) 27/x; polar diameter (10 specimens) 14 (18) 23 ju. Remarks'. The original grain shape as deduced from measurements and grain orientations appears to be subspheroidal to oblate spheroidal. The tetrad arrangement and apertural position have not been deduced although Muller (1970) suggested that the aperture is distal. The aperture is expressed in the nexine as a four or five (rarely three) branched opening, the branches radiating from the ?distal pole and extending towards the equator. The individual branches of the aperture are ± parallel-sided but have enlarged equatorial extremities. The columellate sexine may be entire over the nexine sulcus where it exhibits sculptural modification; often the sexine is partially or completely broken down to expose the sulcus. Comparison: The Australian grains appear to be comparable to those described and illustrated by Hedlund & Norris (1968) and are similar to the specimen figured by Doyle (1969, Plate 1, Fig. i) as Clavatipollenites. Doyle also illustrated (Plate 1, Fig. h) a threebranched form similar to Groot & Groot's species Apiculatisporis vulgaris (1962, Plate 6, Figs 4, 6, 7) and Liliacidites trichotomosulcatus of Singh (1971, Plate 29, Figs 5-7). Distribution: Of persistent occurrence in Cenomanian and Turonian horizons of Bathurst and Melville Islands and with sporadic representation in probable CenomanianTuronian sediments of the Otway Basin. Hedlund & Norris (1968) reported the species from ?late middle Albian of Oklahoma; the similar type figured by Doyle (1969) is from ? Albian (Patapsco Formation) of eastern U.S.A. *Lat. squamosus, scaly.

Subturma PTYCHOTRIPORINES Naumova, 1939 Genus Nyssapollenites Thiergart, 1937 Type species (by original designation): Nyssapollenites pseudocruciatus (Potonie) Thiergart, 1937. Remarks'. Singh (1971) demonstrated that the genus is appropriate for early Upper Cretaceous ± spheroidal, tricolporate pollen having a faintly roughened exine. Comparable features are exhibited by one of the Australian midCretaceous species, Nyssapollenites squamosus sp. nov., here attributed to the genus. The other species, N. lanosus sp. nov., described below is not strictly conformable with the genus since it has weakly developed ora and may be regarded as colporoidate. Affinity: The nyssaceous affinity suggested by Thiergart (1937) and Potonie (1960) for the type species, N. pseudocruciatus (Potonie) Thiergart, 1937, may not necessarily be applicable to the Australian Cenomanian-Turonian grains here referred to the genus. Pollen of similar morphology are known in many extant families, for example in the Araliaceae, Cornaceae, and Marcgraviaceae (see Erdtman, 1952, pp. 60, 130, 264). The last-named family produces colporoidate grains with the apertures arranged according to Fischer's Law, features observed in N. lanosus. Nyssapollenites squamosus* sp. nov. (Plate 5, Figs 9-15) Diagnosis: Pollen grains isopolar, tricolporate, oblate; amb anguloaperturate, triangular with straight to concave sides. Colpi developed in sexine, widest at equator (2-3^), tapering towards polar extremities; in polar view extending i distance to poles, sometimes appearing longer as a result of sexine splitting beyond polar extremities. Ora well defined, circular, 1.5-2^ in diameter, sunk 2-3 jm below sexine surface. Exine 1-1.5^ thick, differentiation into nexine and sexine obscure in all regions of grain except at apertures where sexine is thicker than nexine. Exine surface with low (less than 0.1//, high), broad-based (diameter 0.5^) elevations that are irregularly polygonal in plan view; elevations delineated by narrow grooves and/or minute pits; surface relief reduced at aperture margins. Dimensions'. Equatorial diameter 12 (16) 19JJL; polar diameter (3 specimens) 11 (12) 14/x.

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 19 Holotype: Preparation S301/1, 28.5 109.2, linearly arranged, minute pits. In corroded exY.1480; Plate 5, Figures 9-10. Polar aspect. amples sexine surface is minutely punctate. Amb convexly subtriangular, 16/x in diameter. Dimensions'. Equatorial diameter 12 (15) Colpi extend 5/x from equator to pole; pores 20/*; polar diameter 12 (14) 18/x. 1.5^ in diameter, sunk 2^ below sexine surface. Holotype: Preparation S336/3, 32.4 94.1, Exine 1.5 jx thick, with a distinctly roughened Y.1475. Plate 5, Figures 1-2. Polar aspect. surface (OL pattern). Amb circular, diameter 16/x. Colpi terminatType locality: Bathurst Island, Patinguming 4-5jx from poles; 1.5/x wide at equator; putti; sample Ni 193. Turonian. small ora (1^ diameter) developed in nexine Remarks and comparison: Grains of this at equatorial regions of colpi. Exine 1.5/x thick, species are oblate and are almost invariably surface with irregular elevations 0.2-0.5/x basal orientated in polar view in glycerine jelly diameter. mounts. Examples examined by means of the Type locality: Bathurst Island, Pulliamanscanning electron microscope revealed the equatorial features of the apertures (Plate 5, dera; sample Ni 192. Turonian. Fig. 15). The precise exine differentiation and Remarks and comparison: The colporoidate structure have not been determined; optical nature of the apertures was determined from sections of the exine at magnifications of X observations under the scanning electron and 1,000 under the light microscope appear homo- light microscopes (Plate 5, Fig. 5); in several geneous, except in apertural regions where a other examples the apertures may be more clear differentiation into sexine and nexine is appropriately termed colpate (Plate 5, Fig. 8). This evidence suggests that the species discernible. Nyssapollenites squamosus sp. no v. re- may have apertures transitional between colpi sembles N. albertensis Singh, 1971, Tricolporo- and oriferous colpi. Reference of the grains pollenites triangulus Groot, Penny, & Groot, to Nyssapollenites must thus be considered as 1961, and T. fortis Groot, Penny, & Groot, tentative, despite their obvious morphological 1961, but is distinct in having shorter colpi resemblance to N. squamosus. The latter species is distinguishable in having more clearly and a more conspicuously sculptured exine. Distribution: Recovered from Bathurst defined and sunken ora, shorter sexine colpi, Island in late Cenomanian and Turonian hori- and a convexly triangular amb. zons and from probable Cenomanian-Turonian Nyssapollenites lanosus sp. nov. is similar strata of the Otway Basin. to Tricolporo pollenites aliquantulus Hedlund, 1966 but is larger and has more prominent sculpture. T. distinctus Groot & Penny, 1960 Nyssapollenites lanosus* sp. nov. and T. orbiculatus Groot, Penny, & Groot, (Plate 5, Figs 1-8) 1961 display more clearly defined pores and Diagnosis'. Pollen grains free or united in are described as psilate. tetrahedral tetrads, tricolpate to tricolporoidate, Distribution: The species is represented in isopolar, spherical to subprolate; amb circular Cenomanian and Turonian sediments of to fossaperturate. Apertures arranged accor- Bathurst Island and the Otway Basin. ding to Fischer's Law, colpate to colporoidate; sexine colpi extending almost to poles, ± parallel-sided or widest (2-3^) at equator, Turma POROSES Naumova emend. Potonie, 1960 tapering towards polar extremities. Nexine occasionally with small, circular (1-2/x diaSubturma TRIPORINES Naumova emend. meter), equatorially situated, indistinct oroids; Potonie, 1960 usually nexine is split in apertural regions, the Genus Triorites Cookson ex Couper, 1953 splits conforming in position and shape to Type species (by subsequent designation of sexine colpi. Exine indistinctly differentiated into nexine and sexine; nexine 0.5/* thick; Couper, 1953; p. 60): Triorites magnificus sexine thicker (1-1.5/x), ?thinning at colpal Cookson, 1950. margins. Surface with low, irregular elevations Remarks: The genus is used here in the (less than 0.1/*, high, 0.2-0.7/* basal diameter) sense of Couper (1953) and not in the rethat are delineated by narrow grooves or stricted version of Potonie (1960). This usage

* Lat., lanosus, woolly. Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

20

MARY E. E

follows that of Dettmann & Playford (1968) and Muller (1968) who discussed some of the difficulties associated with this broad generic category. The three forms described and recorded herein are the oldest known representatives of triporate pollen recovered from Australian sediments. Two species, Triorites minor Couper, 1953 and T. punctulatus sp. nov., exhibit little sculptural detail and may be shown eventually to be more appropriately placed in En gelhardtioidites Potonie, Thomson, & Thiergart, 1950. The third type, Triorites sp., has a reticulate sexine and in some respects resembles fossil pollen allocated by Cookson (1950), Couper (1953, 1960), and Harris (1965) to Proteacidites Cookson ex Couper, 1953, which like Triorites is a broad generic category requiring detailed study and revision. Affinity: No precise affinity can be suggested for grains here referred to Triorites. Groups of several specimens (?incomplete tetrads) of T. minor have been observed, and it is of interest that in these the apertural arrangement conforms with Garside's Law. However, complete tetrads must be found before any definite statement on the apertural arrangement is made. A Garside-type arrangement may of course suggest a possible affinity with the Proteaceae or with ancestors of the family. Triorites minor Couper, 1953 (Plate 6, Figs 1-3) Remarks'. Grains referred to Triorites minor by Couper (1953, 1960), Dettmann & Playford (1968), and in the present study have a sexine that is weakly thickened about the small (1-2n), circular ora. The sexine surface exhibits a slight, but perceptible pattern at magnifications of X 1,000; at higher magnifications under the scanning electron microscope, the sexine surface shows low, irregularly-based elevations (0.3-0.5^ basal diameter, 0.1^ high) that are separated by narrow (less than 0.1 /a wide) shallow depressions (Plate 6, Fig. 1). Distribution: Recovered from Turonian of Bathurst Island and probable Turonian of the Otway Basin. Dettmann & Playford (1968, 1969) showed that the species extends into latest Cretaceous sediments of the Otway Basin. Couper (1960) quoted a SenonianUpper Oligocene range for the species in New Zealand. * Lat., punctulum, punctate.

Triorites punctulatus* sp. nov. (Plate 5, Figs 16-21) Diagnosis: Pollen grains triorate, isopolar, oblate, anguloaperturate; amb triangular with straight to weakly convex sides. Ora circular, 2-6/x diameter, developed in both nexine and sexine. Exine two-layered; nexine 0.75^ thick, sexine 0.5-0.7^ thick. Under the light microscope at magnification of X 1,000, sexine appears faintly and finely columellate and shows a faintly microreticulate surface. At higher magnifications under scanning electron microscope, sexine surface has low undulations and minute pits (less than 0.1 ^ in diameter). Dimensions: Equatorial diameter 15 (18) 25fi; polar diameter (3 specimens) 11 (12) 18 fJL. Holotype: Preparation S336/3, 26.1 100.4, Y.1484; Plate 5, Figures 16-17. Polar aspect. Amb triangular with weakly convex sides, diameter 20/x. Exine 1.5^ thick; nexine 0.15^ thick; sexine 0.75fx thick, faintly and finely columellate and with a faint OL surface pattern. Ora 4-6/a diameter. Type locality: Bathurst Island, Pulliamandera; sample Ni 192. Turonian. Comparison: The species resembles Triorites minor, but is distinguishable in having a thicker exine and larger ora which lack thickened rims. T. fragilis Couper, 1953 is considerably larger and has a thinner exine; T. africaensis Jardine & Magloire, 1965 has similar sculpture but differs in its smaller ora and larger size. Distribution: Recovered from Turonian of Bathurst Island and the Otway Basin. Triorites sp. (Plate 6, Figs 4-7) Description: Pollen grains triorate, isopolar, oblate, anguloaperturate; amb triangular with straight to weakly convex sides. Ora circular, 3-4jx diameter. Exine 1.5^ thick; nexine and sexine each 0.75//, thick. Sexine columellate; columellae slender (ca 0.1/x basal diameter), spaced ca 0.1 ^ apart, fused at tips to form imperfect surface reticulum. Sinuous muri 0.4^ wide; lumina irregular, often interconnected, largest in polar regions (2jjl), decreasing to minute pits at equator. Dimensions (6 specimens): Equatorial diameter 20 (22) 25/*. Comparison: Triorites sp. resembles Proteacidites retiformis Couper, 1960 but is dis-

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 21 tinguishable in having a smaller size, smaller The sexine and apertural characters disora, and an imperfect reticulum. tinguish Australopollis obscurus from other Distribution: Recovered from Turonian Australian mid-Cretaceous pollen described in this paper. strata of Bathurst Island. Distribution: Known only from southeastern Subturma POLYPORINES Naumova emend. Australia. In the Otway Basin it appears initiPotonie, 1960 ally in sediments believed to be of Turonian age {cf. Dettmann & Playford, 1968, 1969), Genus Australopollis Krutzsch, 1966 Type species (by original designation): Aus- and it completes its range in the late Palaeocene tralopollis obscurus (Harris) Krutzsch, 1966. (Stover & Evans, 1973; cf. Harris, 1965). Turma JUGATES Erdtman, 1943 Australopollis obscurus (Harris) Krutzsch, Subturma TETRADITES Cookson, 1947 1966 Genus Senectotetradites* gen. nov. (Plate 1, Figs 9-12) Type species (here chosen): Senectotetradites 1965 Stephanoporopollenites obscurus Harris, varireticulatus sp. nov. p. 95; Plate 29, Figures 15-17. 1966 Australopollis obscurus (Harris) Krutzsch, Diagnosis: Pollen grains tricolpate, prolate p. 38. to oblate, united in tetrahedral tetrads or Description: Pollen grains zoniporate, oblate rarely free. Colpi disposed meridionally, in to subspherical; amb circular to subcircular. tetrads arranged according to Fischer's Law colpi meet two and two at six points on Exine 1.5-2/x thick, nexine thicker (1-1.25^) (thetetrad). Exine columellate, the columellae than sexine (0.5-0.7/x) which is columellate. the fused at their distal extremities to form a Columellae fused to form a finely meshed reti- foveolate to reticulate surface pattern. culum (muri ca 0.1/x wide, lumina ca 0.2/x Comparison: Senectotetradites gen. nov. has diameter) that supports a tectum which has densely-packed, fine, low, irregularly based meridionally situated colpate apertures and is conate processes (ca 0.1 p, high, and 0.2^ in thus distinct from the tricolporate grains of basal diameter). Pores five to seven in number, Ericipites Wodehouse, 1933 and the type formed in nexine as subcircular to elliptical species of Dicotetradites Couper, 1953. Gynoapertures, 4-8/x diameter. Sexine continuous cardiidites Bobrowska, Elhai & Erdtman, over pores or with irregular openings formed 1962 has baculate to verrucate sculpture and from partial to complete break-down of sexinal Retitetradites Pierce, 1961 is distally monocolpate. elements in apertural regions. Affinity: Morphologically similar pollen Dimensions'. Equatorial diameter 29 (34) 37^; polar diameter (5 specimens) 22 (24) ('young pollen grains still united in fours', Erdtman, 1952, p. 287) occur in Nelumbo 26/JL. Adams, which on pollen morphologiRemarks and comparison: The sexine struc- (Tourn.) grounds is considered an 'isolated' genus ture of Australopollis obscurus (Harris) cal within the (Erdtman, loc. cit.) Krutzsch, 1966 is considerably more intricate and referredNymphaceae to the Nelumbonaceae by Takhaand complex than other sexine structures ob- tajan (1969). served in Australian mid-Cretaceous angiospermous pollen. The sexine is tectate, the Senectotetradites varireticulatust sp. nov. tectum supported on an infratectal reticulum, and itself supporting supratectal conate pro(Plate 6, Figs 12-16) cesses. The nexinous pores are covered by the Diagnosis: Pollen grains united in tetrasexine which may be irregularly ruptured to hedral tetrads, tricolpate, prolate subspheroidal, expose the pores. Several examples have been or oblate; amb circular. Colpi narrow slits examined by means of the scanning electron which have pointed extremities and ± parallel microscope; in these the number of pores de- sides, length i polar diameter of grain; veloped in the sexine varies from one to three, arranged according to Fischer's Law. Nexine the remaining nexinous pores being covered by 1 ^ thick, appears homogeneous under light an uninterrupted layer of the sexine. microscope. Sexine differentially thickened, * Lat., senectus, old t Lat., varius, variably; and Lat. reticulatus, netted. Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973. 9

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MARY E. DETTMANN

thickest (2.5-3//,) about distal pole, thinning to composed of thin (ca 0.2//,) nexine which 1.5-2.5//, at equator and 0.5-1//, over proximal appears homogeneous under light microscope. surface. Sexine columellate; columellae largest Sexine thicker, thickest (1.5-2//,) about poles, and more widely spaced about distal pole (2-3//, thinning to 1-1.5//, at equator; columellate and high, 0.5//, basal diameter, head diameter, with a foveolate surface pattern. Columellae spaced 1-2//, apart) decreasing in size and 1-2//, high, closely spaced (ca 0.5//, apart), spacing towards equator and on proximal sur- with cylindrical columns (0.5/x basal diameter) face. Columellate heads ± flat-topped, fused and expanded heads (0.75-1//, diameter). Coluto form a surface reticulum which attains mellae fused at heads and occasionally along greatest mesh diameter in mesocolpia. Muri of their length to form a foveolate tectum. reticulum 1-1.5//, wide, flat-topped, enclose Foveolae subcircular to elliptical, 0.3-0.5//, diapolygonal lumina which are as much as 5//, meter, spaced 0.5-1.5//, apart. Columellae often wide in mesocolpia, decreasing in size towards exposed along colpal margins which assume a distal pole and along colpal margins. Surface jagged outline. sculpture of proximal contact areas not known. Dimensions (of individuals in tetrad): EquaDimensions (of individuals in tetrads): torial diameter 23 (27) 33//,; polar diameter Equatorial diameter 19 (27) 33//,; polar dia- 17 (28) 33/z. Holotype: Preparation S340/1, 23.5 90.0, meter 19 (28) 35//,. Holotype: Preparation K340/2, 32.0 89.9, Y.1491; Plate 6, Figure 8. Four tricolpate Y.1495; Plate 6, Figure 13. Tetrahedral tetrad; grains arranged in tetrahedral tetrad with colpi grains tricolpate with colpi arranged according meeting two and two at six points on the tetto Fischer's Law; grain diameter—equatorial rad. Grains subspheroidal, diameter 33//,. Colpi 32/x, polar 32//,. Nexine 1//, thick, sexine 2.5//, with pointed extremities, gaping (3/x) at equaat distal pole, 2//, at equator. Columellae 2/x tor and with jagged margins. Exine 2.5/x thick; high, basal diameter 0.5/x, head diameter 1 //,. sexine 2//, at distal pole thinning to 1.25//, at Surface reticulum with muri 1//, wide, enclos- equator. Columellae 1-2//, high, columns 0.5/x ing polygonal lumina 3//, in diameter in meso- basal diameter, heads 1//, diameter. Heads fused colpal areas and 1//, in diameter about distal to form foveolate surface pattern; foveolae subcircular to elliptical, ca 0.5-1//, diameter, pole. Type locality: Queensland, Delhi-Santos spaced 1-1.5//, apart. Type locality: Bathurst Island, Meadinga, Mornington Island No. 1 well, cuttings from sample Ni 102. Cenomanian. 310-20 ft (94-7m). Cenomanian. Remarks and comparison: Although the Remarks'. Pollen of this species are invariably united in tetrahedral tetrads and the proxi- majority of examples observed are united in mal surficial features have not been determined. tetrahedral tetrads, occasional single grains A study of tetrads under the scanning electron have been recorded (Plate 6, Fig. 9). Senectomicroscope indicates that the grains are closely tetradites fistulosus sp. nov. shows a striking adherent and possibly fused over their proxi- resemblance to, and may be conspecific with, tetrad type 1 of Doyle (1969, Plate 3, Fig. mal contract areas. k). It is readily distinguishable from S. Distribution: Widely distributed in Ceno- j,varireticulatus nov. in its uniformly demanian of Bathurst, Melville, and Mornington veloped foveolatesp.surface Single grains Islands. The species has been observed also in are readily distinguished pattern. from Tricolpites sp. probable Cenomanian strata of the Otway (this paper) and T. cooksonae in having a Basin. thicker exine and a foveolate surface pattern. Senectotetradites fistulosus* sp. nov. Distribution: Recovered from the Ceno(Plate 6, Figs 8-11) manian of Mornington, Bathurst, and Melville Diagnosis: Pollen grains united in tetra- Islands. hedral tetrads or occasionally free, tricolpate, DISTRIBUTION AND SEQUENCE OF subprolate to suboblate; amb circular. Colpi ANGIOSPERMOUS POLLEN FLORAS narrow or irregularly gaping slits, with pointed The mid-Cretaceous pollen types described extremities, length 2/3 polar diameter; meeting two and two at six points on tetrad (i.e. in the systematic section include the oldest arranged according to Fischer's Law). Exine known angiospermous pollen from eastern *Lat., fistulosus, porous.

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN Australia. The pollen occur in assemblages that are dominated by derivatives of bryophytes, lycopods, pteridophytes, and gymnosperms. Dettmann & Playford (1969) noted the uniformity of palynological floras in eastern Australia during the Early Cretaceous. This uniformity is most marked in NeocomianAptian strata, but by the Albian there are some notable microfloral dissimilarities between the north and south of the eastern Australian region. In the north, gleicheniaceous, schizaeaceous, and the possible gymnosperm Hoegisporis are well represented; in southern regions these types are subordinate and are overshadowed by diverse hepatic and lycopod species. These changes possibly reflect contemporaneous alterations in the floras in response to processes of plant migration and evolution. Significantly, angiospermous pollen make their first appearance during this time interval (Albian).

23

Many of the Albian-Turonian pollen species described in this investigation have widespread distribution in eastern Australian deposits. Others, however, show some geographic restriction or their vertical occurrences are seemingly distinct in different regions. A comparison of the angiospermous pollen floras obtained from the principal areas of study is attempted here to gain some appreciation of the geographic limitations of the species and possible migratory trends (Fig. 3). Similarly the eastern Australian angiosperm pollen floras are compared with those from overseas Albian-Turonian sediments. Composition of eastern Australian assemblages The oldest indisputable angiospermous pollen so far recognized in eastern Australia occur at late middle Albian horizons. These earliest records are from basal horizons of the

Fig. 3. Vertical distribution of pollen species within Albian-Turonian strata of Bathurst and Melville Islands and the Great Artesian and Otway Basins. Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

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MARY E. DETTMANN

Allaru Mudstone and equivalents in the Great Artesian Basin, and accord with Burger's (1971) observations. The oldest types recognized are reticulate tricolpate and tricolpoidate forms of Phimopollenites pannosus (Dettmann & Playford) comb. nov. and Rousea georgensis (Brenner) comb. nov. which occur sporadically in middle Albian sediments, becoming more numerous in the late Albian. Here they are joined by other tricolpate species, Tricolpites minutus (Brenner) comb. nov. and Phimopollenites augathallaensis (Burger) comb, nov., and by rare examples of the monosulcate form Clavatipollenites sp. In the Otway Basin tricolpates and tricolpoidates are once again the earliest detected angiospermous pollen. Here they possibly make a somewhat later appearance in the Albian (late Albian). They are less diverse, being represented only by P. pannosus and R. georgensis. Monosulcate grains have not been observed in the late Albian of the Otway Basin. During the Cenomanian, tricolpates became more numerous and diverse in eastern Australia with the Albian forms persisting (Fig. 3). Clavatipollenites is rare, but occurs throughout the region studied, making its initial appearance in the Otway Basin. Asteropollis asteroides Hedlund & Norris, 1968, tricolpate tetrads of Senectotetradites gen. nov., and tricolporoidate to tricolporate forms of Nyssapollenites Thiergart, 1937 were widely and apparently synchronously established. Several species appear to have restricted geographical distribution; these include Liliacidites cf. kaitangataensis Couper, 1953, L. cf. intermedins Couper, 1953, Striatopollis cf. paraneus (Norris) Singh, 1971, Cupuliferoidaepollenites cf. parvulus (Groot & Penny) comb. nov. and Tricolpites cooksonae sp. nov. which are known only from the northern Australian Cenomanian (Bathurst, Melville, and Mornington Islands). During the Turonian many of the Albian and Cenomanian tricolpate forms disappeared and were replaced by forms with welldeveloped porate apertures. The Turonian floras of Bathurst Island and the Otway Basin contain many species in common, for example, Asteropollis asteroides, Clavatipollenites sp., Nyssapollenites lanosus sp. nov., N. squamosus sp. nov., Triorites punctulatus sp. nov., and Triorites minor Couper, 1953 (see Fig. 3 ) . However, there are certain striking differences between the Turonian pollen floras of Bathurst Island and the Otway Basin. Northern species not found in the latter include Liliacidites cf.

intermedins, Triorites sp., and Cupuliferoidaepollenites cf. parvulus together with several undescribed periporate forms. Species restricted to the Otway Basin during this time-interval include Phimopollenites pannosus (which makes its last appearance on Bathurst Island during the Cenomanian) and the periporate form Australopollis obscurus (Harris) Krutzsch, 1966. Australasian comparisons The relationships of eastern Australian midCretaceous angiospermous pollen floras with those of Western Australia are virtually unknown since no detailed information on the latter area is available. Balme (1964, p. 75) stated that 'the oldest definite angiosperm pollens known in Western Australia are of Upper Senonian age', but has subsequently observed (pers. comm.) tricolpate grains in the ?Albian-?Cenomanian Osborne Formation of the Perth Basin. Doyle (1969) in referring to Kemp (1966, unpubl.) alluded to Albian occurrences of Clavatipollenites and tricolpate grains from the northwestern part of Western Australia. Farther north, in Papua, angiospermous pollen assemblages have been recovered from subsurface marine late Albian to Turonian sediments. The pollen floras are poorly preserved but appear to be closely similar to assemblages from Bathurst and Melville Islands (Dettmann, unpublished data). In New Zealand, dicotyledonous grains appeared initially during the Albian (Couper, 1960); the types represented have not been specified, thus precluding comparison with eastern Australian types. Forms similar to, if not identical with, Couper's (1953, 1960) Triorites minor, Liliacidites kaitangataensis, and L. intermedins seemingly appeared earlier in Australia (Cenomanian and Turonian) than in New Zealand (Senonian and later). Extra-Australasian comparisons Angiospermous pollen are widely distributed in Albian and later strata of areas beyond Australasia. There are records, however, of presumed angiospermous grains from significantly older European and North American strata. In these areas Clavatipollenites, a monosulcate form accepted by most authors as of angiospermous origin, appeared as early as the Barremian (Couper, 1958; Kemp, 1968; Doyle, 1969 and see other references cited therein). Other putative and even earlier records of the genus include Jurassic of central Europe, Canada, and Egypt (Schulz, 1967; Pocock,

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 1962; Helal, 1966, and cf. Kemp, 1968). In eastern Australia Clavatipollenites is not known until the late upper Albian (Great Artesian Basin) and Cenomanian (Otway Basin). This evidence may support a relatively slow migration of the genus into the Australian region, possibly by way of the Pacific Basin. Moreover, the morphologically related tri- to pentachotomosulcate form, Asteropollis, appears to have reached Australia at a significantly later date than in North America and Europe. From Oklahoma Hedlund & Norris (1968) reported their A. asteroides from the ?late middle Albian, but this same species does not occur in Australia until the late Cenomanian. Other representatives of Asteropollis occur in the late middle Albian of western Canada (Liliacidites trichotomosulcatus Singh, 1971), AptianAlbian of Portugal (Apiculatisporis vulgaris Groot & Groot, 1962), and mid-Lower Cretaceous of eastern U.S.A. (Doyle, 1969, p. 6). The initial appearance of tricolpate grains in eastern Australia is late middle Albian, an age synchronous with, or slightly younger than, reported occurrences from Europe, U.S.S.R., North and central America, Malaysia, and equatorial Africa (Brenner, 1963, 1967, 1968; Boltenhagen, 1965; Cahoon, 1968; Doyle, 1969; Groot & Groot, 1962; Hedlund & Norris, 1968; Jardine & Magloire, 1965; Kemp, 1968; Muller, 1968; Norris, 1967; Playford, 1971; Singh, 1971; Zaklinskaya, 1960, 1962). However, a significantly earlier occurrence (Berriasian-Valanginian) of tricolpate (or ?tricolporate) grains has been reported from the Netherlands (Burger, 1966); this is accepted by Muller (1970). The earliest occurring Australian tricolpates have a reticulate exine; in Cenomanian and Turonian species the exine may be almost smooth. A parallel situation occurs in eastern U.S.A. (Doyle, 1969). Similarly tricolpate grains permanently united in tetrads (Senectotetradites) seem to have appeared contemporaneously in eastern Australia and eastern U.S.A. (Doyle, 1969). Eastern Australian tricolporoidate and tricolporate pollen are evident only at late Cenomanian horizons, later than reported inceptions in North America (?late Albian and early Cenomanian respectively—see Doyle, 1969; Singh, 1971). Triporate pollen appeared in eastern Australia during the Turonian, again at a probable later date than in other areas, including Malaysia (Cenomanian-Senonian; Muller, 1968), west Africa (upper Albian-lower Ceno-

25

manian; Jardine & Magloire, 1965), North America (late Cenomanian; Doyle, 1969) and Europe (Cenomanian; Groot & Groot, 1962). The Australian triporates have simple pores, exhibiting morphological similarity to possible proteaceous types described from Cenomanian of west Africa (Jardine & Magloire, 1965). The Normapolles type, characteristic of Europe and North America has not been detected in Australasian strata. The unusual tricolpodiorate and polyporate forms described from late Albian-Cenomanian of central America and west Africa (Brenner, 1968; Jardine & Magloire, 1965) are not represented in eastern Australia. Polyporates do occur at Turonian and younger horizons, where forms with ragged pores, including A ustralopollis, are represented. Present data, incomplete as these admittedly are, suggest that Australian mid-Cretaceous assemblages show closest affiliation with those that have been recorded from North America. This similarity is most marked in the midAlbian-Cenomanian tricolpate types, Forms similar to, if not identical with, the following species occur within this time interval in eastern Australia and North America: Tricolpites minutus, Rousea georgensis, Striatopollis paraneus, Cupuliferoidaepollenites parvulus, and Senectotetradites fistulosus (see Brenner, 1963; Norris, 1967; Singh, 1971; Playford, 1971; Groot & Penny, 1960; Doyle, 1969). Australian late Cenomanian-Turonian tricolporate and tricolporoidate species, Nyssapollenites squamosus and N. lanosus, exhibit some resemblance to middle-upper Cenomanian types represented in North America. EVOLUTIONARY IMPLICATIONS The Australian mid-Cretaceous angiospermous pollen described herein, although morphologically simple in comparison with Early Tertiary to Recent angiosperm pollen, display a wide range of apertural characters and exine sculpture and structure. These pollen morphological characters may be evaluated against a chronological time-scale to establish evolutionary trends within the Angiospermae. Syntheses of morphological characters in this context have been attempted recently, not only for fossil angiospermous pollen grains, but also for spores (s. str.)9 pre-pollen, and gymnospermous pollen (Doyle, 1969; Muller, 1970; Chaloner, 1970). The validity of using pollen morphology in evolutionary and phylogenetic studies hinges, at least partially, on the premise

Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

MARY E. DETTMANN

26

that pollen morphologies 'are expressions of the genotype of the haploid spore, and not something impressed upon the wall by the environment of the anther' (Heslop-Harrison, 1963, pp 22-3). This hypothesis is still under debate (Heslop-Harrison, 1968), and consequently phylogenetic conclusions based upon pollen morphology alone must be viewed with caution. Nevertheless, pollen morphology has found significant application in plant taxonomy (Erdtman, 1952, 1969) and has undoubted potential in contributing to our knowledge of the history of the angiosperms. The most striking evolutionary trends seen in the early angiospermous pollen are in the shape and position of the apertures. Doyle (1969, Fig. 6) has documented an evolutionary sequence based upon initial appearances of distinct apertural characters. Thus, in recognizing that the heteropolar, monosulcate form Clavatipollenites preceded isopolar, tricolpate types, Doyle (1969) suggested derivation

of tricolpate apertures from trichotomosulcate variants (Asteropollis) of Clavatipollenites. If such differentiation did take place, then we can assume from evidence documented herein (i.e. the appearance within Australia of tricolpate forms prior to the incoming of Clavatipollenites and Asteropollis) that Australia was not the site of this differentiation. In any case there are arguments against such a derivation. If, as noted by Chaloner (1970), the trichotomosulcus of Asteropollis is disposed radially on the distal surface, then reduction and migration of the sulcus arms would provide a tricolpate type with the apertures arranged according to Garside's Law. Evidence obtained from Australian midCretaceous tricolpates (Table I) indicates that the apertures are arranged instead according to Fischer's Law. Arguments for and against the derivation of tricolpates from forms of the Clavatipollenites-type are based on the assumption that

TABLE I

Tetrad types and apertural configurations and positions in eastern Australian mid-Cretaceous angiospermous pollen species Species

Tetrad type

Clavatipollenites sp. Liliacidites cf. kaitangataensis L. cf. intermedius Asteropollis asteroides

not known not known not known not known

Phimopollenites pannosus P. augathallaensis Tricolpites minutus T. cooksonae T. sp. Striatopollis cf. paraneus Rousea georgensis Cupuliferoidaepollenites cf. parvulus Nyssapollenites lanosus N. squamosus Triorites minor T. punctulatus T. sp. Australopollis obscurus Senectotetradites varireticulatus S. fistulosus

tetrahedral not known tetrahedral tetrahedral not known not known not known tetrahedral tetrahedral tetrahedral ?tetrahedral not known not known not known tetrahedral tetrahedral

Apertural configuration monosulcate monosulcate monosulcate tri- to pentachotomosulcate tricolpoidate tricolpoidate tricolpate tricolpate tricolpate tricolpate tricolpate tricolpate tricolporoidate tricolporate triporate triporate triporate polyporate tricolpate tricolpate

Apertural position not known (? distal) not known (? distal) not known (? distal) not known (? distal) meridional (Fischer's Law) meridional meridional (Fischer's Law) meridional (Fischer's Law) meridional meridional meridional meridional (Fischer's Law) meridional (Fischer's Law) meridional (Fischer's Law) meridional (?Garside's Law) meridional meridional meridional meridional (Fischer's Law) meridional (Fischer's Law)

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN 27 the sulcus in the latter type (and the mor- (Jardine & Magloire, 1965; Brenner, 1968), phologically related Asteropollis) is distal. but these Albian types have the pores arranged Moreover, it is assumed that the arms of the globally. In Australopollis the pores are trichotomosulcus of Asteropollis are disposed meridional, and in some features (apertures radially in a tetrahedral tetrad. These in- with irregular outline and opercula) the genus ferences are based upon comparisons with is similar to forms of the Stephanocolpites living mono- and trichotomosulcate grains and fredericksburgensis-typQ. need to be substantiated from fossil evidence. Trends in sculptural and shape features of Mention is made here of the remarkable grains in Australian material closely parallel morphological similarity in character of aper- evidence documented by Doyle (1969) from ture (irregular to elliptical colpoids with oper- eastern U.S.A. In Australia the earliest tricula) and sexine structure/sculpture seen in aperturate forms are prolate and have reticuClavatipollenites, Asteropollis asteroides, late sculpture; by Turonian times these were Stephanocolpites fredericksburgensis Hedlund largely replaced by oblate forms with the & Norris, 1968, and Phimopollenites. These columellae more completely fused to form forms could be arranged in a morphological psilate to minutely pitted surfaces. Complete series, although it is difficult to envisage deri- fusion of the columellae is seen in the polyvation of a tricolpoidate Phimopollenites from porate Australopollis (Turonian) which has a the tetra- to pentacolpoidate Stephanopollenites tectate sexine, the tectum supporting supratecfredericksburgensis. Nevertheless, a few ex- tal sculptural processes. The trend towards amples of tetracolpoidate Phimopollenites (P. more complex sexine sculpture/structure is also augathallaensis) have been recorded (Plate 3, recognizable amongst monosulcate grains. The Fig. 16) in the Australian material. In this earlier appearing Clavatipollenites has evenly connexion it is obviously important to establish spaced columellae, whereas in Cenomanian accurately the geographical distribution and species of Liliacidites the columellae are of initial appearance of Phimopollenites. The pos- differential development and spacing. tulated derivation of Phimopollenites is not intended to be applicable to all tricolpate forms; PHYLOGENETIC IMPLICATIONS indeed, there is evidence from North America As noted in the systematic section the genand Europe that the earliest tricolpates (of the Tricolpites-type) appeared before the incep- eric or even familial affinities of the Australian tion of Asteropollis asteroides and Stephano- mid-Cretaceous angiospermous pollen types are largely unknown. Resemblances of certain colpites fredericksburgensis. fossil types to pollen of extant taxa Derivation of tricolporates from tricolpates Australian have been mentioned but these are not inis fully supported by the Australian evidence, tended to imply established affinity. Neverboth in representation of intermediate mor- theless, the pollenan record from Ausphological forms and in sequential appearance. tralian Albian sequences obtained supports evidence Similarly, the appearance of tricolpate forms documented by Doyle (1969) and Muller permanently united in tetrads supersedes the (1970) that the earliest occurring tricolpate introduction of free tricolpates (Doyle, 1969). types probably derived from one or several It seems unlikely that triporates were derived families within the 'primitive' Orders Hamafrom tricolporates within Australia. Here the melidales and Trochodendrales as treated by apertures of the tricolporates are convincingly Cronquist (1968) and Takhatajan (1969). arranged in a Fischer-type tetrad, whereas there During the Cenomanian the tricolpates is some, admittedly meagre, evidence for a diversified and morphological similarities can Gar side-type apertural arrangement of the be detected in several other orders, for example triporates (Table I). As has been discussed the Fabales and Nelumbonales. The late Cenopreviously, Australian (Turonian) triporates manian Turonian tricolporates suggest even exhibit some similarity to the possibly pro- higher orders such as the Cornales. Derivation teaceous-type grains (Triorites africaensis) re- of Australian Turonian triporates is uncertain, corded from the west African Cenomanian; the although they bear some resemblance to Late morphological derivation of this type of grain Cretaceous forms of possible protealean affihas not yet been established. nity. The derivation of the polyporate AustraloCouper (1958), Doyle (1969), and others pollis is uncertain. Polyporate forms are known noted the morphological resemblance of monofrom the Albian of west Africa and Peru sulcate and tri- to pentachotomosulcate pollen Spec.Publs geol.Soc.Aust., 4: pp. 3-34, Pis 1-6, 1973.

28

MARY E. DETTMANN

of Clavatipollenites and Asteropollis to pollen of modern representatives of the dicotyledonous Chloranthaceae, Myristicaceae, and Canellaceae; morphologically similar pollen are also known amongst the monocotyledons, for example in some palms (see Erdtman, 1952, 1969). Other monosulcate forms (Liliacidites) represented in Australian CenomanianTuronian strata are more convincingly of monocotyledonous origin since they resemble closely extant pollen of the Liliaceae and Amaryllidaceae. PALAEOECOLOGICAL CONSIDERATIONS A study of the distribution pattern of dispersed land-derived spores and pollen grains in the light of the adduced depositional environment of the enclosing sediments may provide important data relating to the growth sites of the parent plants. Detailed environmental analyses are outside the scope of this investigation but some observations which may be relevant to this subject are presented below. In the Great Artesian Basin, angiospermous grains appear initially in low frequencies in late middle Albian strata (Allaru Mudstone and equivalents) of all localities studied, from Oodnadatta in the south to Mornington Island in the north. These strata are all of marine origin (Day, 1969). The latest Albian-?early Cenomanian horizons studied from the Eromanga Basin were deposited in lagoonal situations and contain an abundance of angiosperm pollen (up to 60 per cent of total spore-pollen assemblage). In contrast marine late Albianearly Cenomanian strata in Mornington Island No. 1 well yielded low percentages (less than 2 per cent) of angiospermous pollen. A similar situation occurs in the Otway Basin late Albian sequence. Here non-marine sediments deposited along the margins of the basin contain high percentages (up to 60 per cent) of angiospermous grains, but in approximately coeval sediments deposited under marine influences, angiospermous pollen are rare to absent. Thus, during the late middle Albian to early Cenomanian in eastern Australia there appears to be a direct relationship between depositional environment of sediment and numerical abundance of angiospermous grains. The high abundance of angiosperm pollen in near-shore sediments and low percentage in off-shore marine deposits suggest that the pollen were derived from plant communities situated close to the shoreline and that the pollen

was not circulated to any great extent by wind a n d / o r water currents. Palaeotemperature data based upon marine faunas indicate that temperate conditions prevailed in the Great Artesian Basin during the Albian (Day, 1969). This evidence implies that the vegetation inhabiting lowland fringes of the area constituted a temperate flora. Moreover, it has been postulated (Irving, 1964) that during this time-interval northwestern parts of the continent were closer (45-50°S) to the tropics than southeastern regions (7075°S); i.e. that climatic disparity existed between northern and southern regions, with warmer conditions prevailing to the north. Significantly Albian and early Cenomanian angiospermous pollen floras from northern areas (Great Artesian Basin) contain a greater diversity of species than those from the Otway Basin in southeastern Australia. Late Cenomanian and Turonian sequences studied from Bathurst and Melville Islands are wholly marine. Angiospermous grains occur persistently but infrequently (less than 5 per cent) at Cenomanian horizons and in higher proportions (up to 10 per cent) in Turonian samples. In the Otway Basin Cenomanian-Turonian horizons examined were deposited under turbid marine influences (Taylor, 1964), and almost invariably contain angiospermous pollen in proportions of 5-10 per cent. The more consistent occurrence of angiospermous grains at late Cenomanian and Turonian horizons possibly indicates that the angiospermous flora had expanded not only in areal distribution, but also into more varied ecological situations. Climatic evidence from late CenomanianTuronian of eastern Australia is inconclusive. A Turonian ammonite from the Otway Basin yielded a palaeotemperature of 28°C (Dorman, 1966), but associated foraminiferal assemblages were thought by Taylor (1964) to represent a cold water fauna. Cool Turonian conditions in the Otway Basin were suggested by Dettmann & Playford (1969) on the basis of a decrease in number and diversity of schizaeaceous spore derivatives and the introduction of gymnospermous pollen resembling that of Dacrydium franklinii Hook. These two forms probably derived from distinct plant associations within the area, the former from fringe-floras and the latter from upland communities. Similarly, as suggested above, the contemporaneous angiosperms may have been segregated into distinct vegetational belts, the

ANGIOSPERMOUS POLLEN FROM ALBIAN TO TURONIAN

distributions of which were climatically (altitudinally) controlled. No palaeotemperature data are available for the Cenomanian and Turonian of Bathurst and Melville Islands, although Wright (1963) postulated that the ammonite faunas have most affinities with Indo-African and Pacific ones. CONCLUSION Evidence obtained from this study lends some support to the view of Scott, Barghoorn & Leopold (1960), Hughes (1961), Doyle (1969), and Muller (1970) that the primary radiation of angiosperms was initiated during the Early Cretaceous. The conclusions of these authors are based partially upon the oldest occurrences of undisputable angiospermous pollen. However, as Muller (1970) has emphasized, pollen of earliest members of the group may not be represented in the fossil record or may be indistinguishable from gymnospermous pollen. The first appearance of angiospermous pollen in eastern Australia has been dated as late middle Albian, and thus at a significantly later date than initial appearances (BarremianAptian) in other parts of the world, notably Europe and the Americas. This evidence argues against an origin of the Angiospermae in eastern Australia (cf. Takhatajan, 1969) despite the substantial representation of 'primitive' taxa in its present day flora. The introduction of certain distinct morphological types within eastern Australian mid-Cretaceous pollen floras is seemingly later than in other areas of the world. Added to this, Australian pollen assemblages of that age are notably less diverse

29

than those of the Americas and west Africa. These data may point to migrations into, rather than evolution within, the eastern Australian mid-Cretaceous floras. The pollen record suggests that Albian and Cenomanian floras of eastern Australia were fairly uniform in composition (but with greater diversity to the north) and that initial geographic differentiation took place between floras in high and low latitudes during the Turonian. ACKNOWLEDGMENTS This research has been materially aided by the valued co-operation and assistance of a number of institutions and individuals, to all of whom I am sincerely grateful. The Geological Survey of Victoria, Geological Survey of Queensland, Department of Mines of South Australia, Frome-Broken Hill Pty Ltd, and Shell Development (Australia) Pty Ltd have provided samples. I am especially grateful to Mr J. V. Hardy and other members of the Electron Microscope Unit, University of Queensland, for facilitating the study of pollen types by means of the Cambridge Stereoscan IIA microscope. Dr H. A. Leffingwell, Union Oil Company of California, Brea, kindly communicated his method of preparing pollen grains for scanning electron microscope study and sent suitable substrate adhesives. Special thanks are extended to Dr H. T. Clifford and Dr G. Playford of the Departments of Botany and Geology respectively of the University of Queensland, for helpful advice and assistance; and to Mrs E. Burdin (University of Queensland) for drafting the text illustrations.

REFERENCES

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the Australian region. Aust. J. Bot., 8, pp. 75-211. BURGER, D., 1966: Palynology of uppermost Jurassic and lowermost Cretaceous strata in the eastern Netherlands. Leid. geol. Meded 35, pp. 211-276. , 1971: Early Cretaceous angiospermous pollen grains from Queensland. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., 116, pp. 1-10. CAHOON, E. J., 1968: Palynology of the Inyan Kara Group of South Dakota. Am. J. Bot., 55, pp. 724-725. CHALONER, W. G., 1970: T h e evolution of mio-

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30

MARY E. DETTMANN

DORMAN, F . H . , 1 9 6 6 : Australian Tertiary paleo1967: A n temperatures. J. Geol., 74, pp. 4 9 - 6 1 . investigation of microplankton assemblages from the Chalk of the Isle of Wight, Eng- DOUGLAS, J . G., 1 9 6 1 : Microplankton of the Deland. Verh. K. ned. Akad. Wet., 24. flandreidae group in Western District sediCOOKSON, I. C., 1950: Fossil pollen grains of proments. Min geol. J., 6, pp. 1 7 - 3 2 . teaceous type from Tertiary deposits in Aus- DOYLE, J . A., 1 9 6 9 : Cretaceous angiosperm pollen tralia. Aust. J. scient. Res., Ser. B, 3, pp. 166of the Atlantic coastal plain and its evolu177. tionary significance. J. Arnold Arbor., 50, , 1961: H<Pegisporis, a new Australian Crepp. 1 - 3 5 . taceous form genus. Palaeontology, 3, pp. ERDTMAN, G., 1952: Pollen Morphology and Plant 485-486. Taxonomy. Angiosperms. Waltham, Mass. , & BALME, B . E., 1962: Amosopollis cruci, 1969: Handbook of Palynology. An Introformis gen. et sp. nov., a pollen tetrad from duction to the Study of Pollen Grains and the Cretaceous of Western Australia. J. Proc. Spores. Munksgaard, Copenhagen. R. Soc. West. Aust., 45, pp. 97-99. EVANS, P. R., 1966: Mesozoic stratigraphic paly, & DETTMANN, M . E. 1959: Microfloras in nology of the Otway Basin. Rec. Bur. Miner. bore cores from Alberton West, Victoria. Resour. Geol. Geophys. Aust., 1966/69 Proc. R. Soc. Vict., 71, pp. 31-38. [unpublished]. , & EISENACK, A., 1958: Microplankton FAEGRI, K . , & IVERSEN, J . , 1 9 5 0 : Text-book of from Australian and New Guinea Upper Modern Pollen Analysis. Munksgaard, CopenMesozoic sediments. Proc. R. Soc. Vict., 70, hagen. pp. 19-79. , & , 1968: Microplankton from two FREYTAG, I . B., 1966: Proposed rock units for marine Lower Cretaceous sediments in the samples from Gingin Brook No. 4 borehole, Oodnadatta region of the Great Artesian Western Australia. J. Proc. R. Soc. West. Basin. Q. geol. Notes geol. Surv. S. Aust., 18, Aust., 51, pp. 110-122. pp. 3-7. COUPER, R. A., 1953: Upper Mesozoic and CainoGERMERAAD, J . H . , HOPPING, C . A . , & MULLER, zoic spores and pollen grains from New J., 1968: Palynology of Tertiary sediments Zealand. Palaeont. Bull., Wellington, 22. from tropical areas. Rev. Palaeobot. Palyn., , 1958: British Mesozoic microspores and 6, pp. 189-348. pollen grains, a systematic and stratigraphic GROOT, J. J. & GROOT, C. R., 1962: Plant microstudy. Palaeontographica, 103B, pp. 75-179. fossils from Aptian, Albian and Cenomanian , 1960: New Zealand Mesozoic and Cainodeposits of Portugal. Commungoes Servs geol. zoic plant microfossils. Palaeont. Bull., Port., 46, pp. 133-175. Wellington, 32. CRONQUIST, A., 1968: The Evolution and Classi- GROOT, J . J . , & PENNY, J . S., 1 9 6 0 : Plant microfossils and age of nonmarine Cretaceous sedification of Flowering Plants. Houghton ments of Maryland and Delaware. MicroMifflin Co., Boston. paleontology, 6, pp. 2 2 5 - 2 3 6 . DAVEY, R . J . , 1 9 7 0 : Non-calcareous microplankton from the Cenomanian of England, northern GROOT, J . J . , PENNY, J . S., & GROOT, C . R . 1 9 6 1 : Plant microfossils and age of the Raritan, France and North America. Part II. Bull. Tuscaloosa, and Magothy formations of the Br. Mus. Nat. Hist. (Geol.) 18, pp. 3 3 5 - 3 9 7 . eastern United States. Palaeontographica, DAY, R. W., 1964: Stratigraphy of the Roma108B, p p . 1 2 1 - 1 4 0 . Wallumbilla area. Pubis geol. Surv. Qd, 318. H A M M E N , T . VAN DER, 1 9 5 6 : A palynological , 1969: The Lower Cretaceous of the Great systematic nomenclature. Boln geol., Bogota, Artesian Basin; in Campbell, K. S. W. (Ed.), 4, p p . 6 3 - 1 0 1 . Stratigraphy and Palaeontology, Essays in , & WIJMSTRA, T. A., 1964: A palynological Honour of Dorothy Hill, pp. 140-173. Aust. study on the Tertiary and Upper Cretaceous Nat. Univ. Press, Canberra. of British Guiana. Leid. geol. Meded., 30, DETTMANN, M. E., 1963: Upper Mesozoic micropp. 183-241. floras from south-eastern Australia. Proc. R. HARRIS, W . K . , 1 9 6 5 : Basal Tertiary microfloras Soc. Vict., 77, pp. 1-148. from the Princetown area, Victoria, Aus, & PLAYFORD, G., 1968: T a x o n o m y of tralia. Palaeontographica, 115B, pp. 7 5 - 1 0 6 . some Cretaceous spores and pollen grains from eastern Australia. Proc. R. Soc. Vict., HEDLUND, R . W . , & NORRIS, G . , 1 9 6 8 : Spores and pollen grains from Fredericksburgian 81, pp. 69-93. (Albian) strata, Marshall County, Oklahoma. , & , 1969: Palynology of the AusPollen Spores, 10, pp. 129-159. tralian Cretaceous: a review; in Campbell, K. S. W. (Ed.), Stratigraphy and Palaeon- HELAL, A. H., 1966: Jurassic plant microfossils from subsurface of Kharga Oasis, Western tology, Essays in Honour of Dorothy Hill, Desert, Egypt. Palaeontographica, 117B, pp. pp. 174-210. Aust. Nat. Univ. Press, Can83-98. berra. CLARKE, R . F . A . , & VERDIER, J . P . ,

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31 R., I 9 6 0 : Synopsis der Gattungen der Sporae dispersae. III. Teil: Nachtrage Sporites, Fortsetzung Pollenites. Mit Generalregister zu Teil I-III. Beih. geol. Jb., 23. , 1966: Idem. IV. Teil: Nachtrage zu alien Gruppen (Turmae). Beih. geol. Jb., 72. , 1970: Idem. V. Teil: Nachtrage zu alien Gruppen (Turmae). Beih. geol. Jb., 87. , 1971: The relation of Arts. 41 & 42 (1966) to the Palaeobotanical Arts. PB3 & 6 (1952). Taxon, 20, pp. 381-382.

J . , 1 9 6 3 : An ultrastructural study of pollen wall ontogeny in Silene pendula. Grana palynol., 4, pp. 7-24. , 1968: Pollen wall development. Science, 161, pp. 230-237. HUGHES, N . F., 1 9 6 1 : Fossil evidence and angiosperm ancestry. Sci. Prog., London, 49, HESLOP-HARRISON,

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IRVING, E., 1964: Paleomagnetism and its Application to Geological and Geophysical Problems. Wiley, New York. JARDINE, S. & MAGLOIRE, L . , 1 9 6 5 : Palynologie et stratigraphie du Cretace des bassins du Senegal et de Cote d'lvoire. Mem. Bur. Rech. Geol. Min., 49, pp. 1 8 7 - 2 4 5 . KEMP, E. M., 1966: Aptian and Albian miospores from southern England. Ph.D. thesis, Univ. Cambridge [unpublished, not seen]. , 1968: Probable angiosperm pollen from British Barremian to Albian strata. Palaeontology, 11, pp. 421-434. KRUTZSCH, W . , 1 9 5 9 : Einige neue Formgattungen und -arten von Sporen und Pollen aus der mitteleuropaischen Oberkreide und dem Tertian Palaeontographica, 105B, pp. 1 2 5 - 1 5 7 . , 1966: Zur Kenntnis der praequartaren periporaten Pollenformen. Geologie, 15, pp. 16-71. LEFFINGWELL, H . A . , LARSON, D . A . , & VALENCIA,

M. J., 1970: A study of the fossil pollen Wodehouseia spinata. Bull. Can. Petrol. Geol., 18, pp. 238-262. LESLIE, R. B., 1966: Petroleum exploration in the Otway Basin. Proc. 8th Commonw. Min. Metall. Congr., 5, pp. 203-216. LUDBROOK, N. H., 1966: Cretaceous biostratigraphy of the Great Artesian Basin in South Australia. Bull. geol. Surv. S. Aust., 36. MULLER, J . , 1968: Palynology of the Pedawan and Plateau Sandstone Formations (CretaceousEocene) in Sarawak, Malaysia. Micropaleontology, 14, pp. 1-37. , 1970: Palynological evidence on early differentiation of angiosperms. Biol. Rev., 45, pp. 417-450. NORRIS, G., 1967: Spores and pollen from the Lower Colorado Group (Albian-?Cenomanian) of central Alberta. Palaeontographica, 120B, pp. 72-115. PIERCE, R. L., 1961: Lower Upper Cretaceous plant microfossils from Minnesota. Bull. Minn. Geol. Surv., 42. PLAYFORD, G., 1971: Palynology of Lower Cretaceous (Swan River) strata of Saskatchewan and Manitoba. Palaeontology, 14, pp. 533565. POCOCK, S. A. J., 1962: Microfloral analysis and age determination of strata at the JurassicCretaceous boundary in the western Canada plains. Palaeontographica, 111B, pp. 1-95.

, THOMSON, P . W . , & THIERGART, F . ,

1950:

Zur Nomenklatur und Klassifikation der neogenen Sporomorphae (Pollen und Sporen). Geol. Jb., 65, pp. 35-70. REYNOLDS, M . HAWKINS,

A.,

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26-33.

ROUSE, G. E., 1962: Plant microfossils from the Burrard Formation of western British Columbia. Micropaleontology, 8, pp. 187-218. SCHULZ, E., 1967: Sporenpalaontologische Untersuchungen ratoliassischer Schichten im Zentralteil des Germanischen Beckens. Palaont. Abh. Abt. B, 2, pp. 541-633. SCOTT,

R.

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BARGHOORN,

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LEOPOLD,

E. B., 1960: How old are the angiosperms? Am. J. Sci., 258A, pp. 284-299. SINGH, C., 1971: Lower Cretaceous microfloras of the Peace River area, northwestern Alberta. Bull. Res. Coun. Alberta, 28. SKWARKO, S . K., 1966: Cretaceous stratigraphy and palaeontology of the Northern Territory. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., 73. SRIVASTAVA, S. K., 1969: Some angiosperm pollen from the Edmonton Formation (Maestrichtian), Alberta, Canada; in Santapau et al. (Eds), J. Sen Memorial Volume, pp. 47-67. J. Sen Memorial Committee and Botanical Society of Bengal, Calcutta. STOVER, L . E . , & EVANS, P . R . , 1 9 7 3 : U p p e r

Cre-

taceous-Eocene spore-pollen zonation, offshore Gippsland Basin, Australia. Spec. Pubis geol. Soc. Aust., 4, pp. 55-72. TAKHATAJAN, A., 1969: Flowering Plants, Origin and Dispersal. Oliver & Boyd, Edinburgh. TAYLOR, D. J., 1964: Foraminifera and the stratigraphy of the western Victorian Cretaceous sediments. Proc. R. Soc. Vict., 77, pp. 535603. TERPSTRA,

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1962:

Palaeontological examination of samples from Delhi-Santos Mornington Island No. 1 well, Carpentaria Basin, Queensland. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1962/177 [unpublished]. THIERGART, F., 1937: Die Pollenflora der niederlausitzer Braunkohle. Jb. preuss. geol. Landesanst. BergAkad., 58, pp. 282-351.

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MARY E. DETTMANN taceous and Paleogene. Dokl. Akad. Nauk S.S.S.R., 133, pp. 431-434 (in Russian). —, 1962: Importance of angiosperm pollen 1967: Revision of the nomenclature of the for the stratigraphy of Upper Cretaceous and Rolling Downs Group in the Eromanga and Lower Paleogene deposits and botanicalSurat Basins. Qd Govt Min. J., 68, pp. 144geographical provinces at the boundary be151. tween the Cretaceous and Tertiary systems; WRIGHT, C. W . , 1963: Cretaceous ammonites from in Reports of Soviet Palynologists for 1st Bathurst Island, northern Australia. PalaeonInternational Conference on Paly nology tology, 6, pp. 597-614. (Tucson, U.S.A.), pp. 105-113. Akad. Nauk ZAKLINSKAYA, E. D., 1960: The role of angiosperm S.S.S.R., Moscow, (in Russian). pollen in the stratigraphy of the Upper Cre32

VINE, R . R., DAY, R . W . , MILLIGAN, E . N . , CASEY, D . J., GALLOWAY, M . C., & EXON, N . F.,

Mary E. Dettmann, Department of Geology and Mineralogy, University of Queensland, St Lucia, Queensland 4067.

EXPLANATION OF PLATES PLATE 1

All figures X 1,000 unless otherwise specified; from unretouched negatives. Figs 1-5. Liliacidites cf. kaitangataensis Couper, 1953. 1, Lateral view showing ?distal sulcus and rupture on ?proximal surface, Bathurst Island, sample Ni 125, prep. S254/1 20.2 98.9, Y.1430. 2, Polar view, Bathurst Island, Meadinga, sample Ni 98, prep. S339/2 29.0 111.1, Y.1431. 3, 4, High and low foci of sexine sculpture, Bathurst Island, Poupanderie, sample Ni 12, prep. S274/1 22.1 102.0, Y.1432. 5, Scanning electron micrograph, polar view, Bathurst Island, Meadinga, sample Ni 102, prep. S340, specimen not recovered. Figs 6-8. Liliacidites cf. intermedius Couper, 1953. 6, 7, Polar aspect, high and low foci showing ?distal sulcus and rupture about ?proximal polar area, Bathurst Island, Moonkinu, sample Ni 130, prep. S344/1 14.9 95.6, Y.1433. 8, Lateral view, Bathurst Island, Moonkinu, sample Ni 144, prep. S287/1 17.5 92.6, Y.1434. Figs 9-12. Australopollis obscurus (Harris) Krutzsch, 1966. 9, 10, Lateral aspect, high and median foci, Victoria, Port Campbell No. 2 well, core 7, 7,913-30 ft (2,41 l-7m), prep. F082/2 33.3 111.0, Y.1435. 11, 12, Scanning electron micrographs X 3,000 and X 10,000 respectively, lateral aspect showing sexine and apertural detail, Victoria, Flaxmans No. 1 well, core 5, 4,479-96 ft (l,365-71m), prep. F119/5 24.3 96.8, Y.1436. PLATE 2

All figures X 1,000 unless otherwise specified; from unretouched negatives. Figs 1-7. Asteropollis asteroides Hedlund & Norris, 1968. 1, 2, Scanning electron micrographs X 2,500 and X 5,000 respectively of specimen situated in polar aspect, showing detail of sexine in equatorial and ?distal polar regions, Bathurst Island, Meadinga, sample Ni 102, prep. S340/14 21.2 99.5, Y.1437. 3, Polar view, Bathurst Island, Moonkinu, sample Ni 145, prep. S288/1 21.0 102.2, Y.1438. 4, Off-polar aspect, Bathurst Island, Toyungimpi, sample Ni 36, prep. S219b/2 16.9 93.5, Y.1439. 5, Lateral view, loc. as fig. 3, prep. S288/2 20.8 104.1, Y.1440. 6, Oblique view, median focus, Bathurst Island, Pouplimanderie, sample Ni 58, prep. S269/2 25.6 91.8, Y.1441. 7, Lateral view of specimen with partially removed sexine, Bathurst Island, Meadinga, sample Ni 102, prep. S340/2 21.1 103.3, Y.1442.

ANGIOSPERMOUS POLLEN FROM ALBIAN T O TURONIAN Figs 8-10.

ClavatipolIenites sp. 8, Polar view, Queensland, Mornington Island No. 1 well core 1, 459 ft (141.5m), prep. K343/2 21.8 102.1, Y.1443. 9, 10, Polar view high and low foci, South Australia, Oodnadatta No. 1 well, 87 ft (26.4m), prep' D362/1 18.9 108.9, Y.1444.

Figs

11-15. Cupuliferoidaepollenites cf. parvulus (Groot & Penny) comb. nov. 11, 12, Polar aspect, high and low foci, Bathurst Island, Meadinga, sample Ni 102, prep. S340/1 30.2 97.4, Y.1445. 13, Polar view, Bathurst Island, Moonkinu, sample Ni 135 prep. S263/1 13.4 99.2, Y.1446. 14, Tetrad X 500, Bathurst Island, Patingumputti, sample Ni 193, prep. S301/1, 14.9 93.5, Y.1447. 15, Lateral view Queensland, Mornington Island No. 1, core 1, 450 ft (136.8m), prep. K342/3 24.5 104.5, Y.1448.

Figs

16-17. Rousea georgensis (Brenner) comb. nov. Lateral view, high and median foci Bathurst Island, sample Ni 122, prep. S286/2 17.5 90.9, Y.1449.

Figs 18-22. Striatopollis cf. paraneus (Norris) Singh, 1971. 18, Polar view, Bathurst Island, Meadinga, sample Ni 102, prep. S340/2 25.9 101.6, Y.1450. 19. Lateral view Queensland, Mornington Island No. 1 well, cuttings, 400-10 ft (121-4m), prep K341/1 23.8 103.8, Y.1451. 20, Lateral view, loc. as fig. 18, prep. S340/2 14.9 91.4, Y.1452. 21, 22, Scanning electron micrographs X 3,000 and X 11,500 respectively of specimen situated in off-polar aspect, showing sculptural and apertural detail, Bathurst Island, loc. as fig. 18, prep. S340/14 19 1 99 0 Y.1453. PLATE 3

All figures X Figs

1-11.

1,000 unless otherwise specified; from unretouched negatives.

Phimopollenites pannosus (Dettmann & Playford) comb. nov. 1-6, All from South Australia, Haddon Downs No. 5 bore, 465 ft (141.4m); 1, 2, holotype high and median foci, Y.353 (see Dettmann & Playford, 1968; pi. 8, fig. 6)' 3, 4, lateral view, high and median foci, Y.355 (see Dettmann & Playford, 1968; pi. 8, fig. 8 ) , 5, polar view, high focus showing operculoid membranes in colpal regions, prep D 3 5 9 / 2 28.0 88.9, Y.1454. 6, polar view, median focus showing partially broken down operculoid membranes, prep. D359/1 26.1 82.3 Y 1455 7, Off-lateral aspect, median focus showing elliptical colpoids South Australia Oodnadatta No. 1 well, 87 ft (26.4m), prep. D362/1 15.3 89.2, Y.1456! 8, 9, Scanning electron micrographs of specimen in polar and lateral aspect, X 3,000, loc. as figs 1, 2, prep. D351/4 21.1 96.0, Y.1457. 10, 11, Scanning electron micrographs X 20,000 of specimens from loc. as figs 1, 2, showing sexine detail in polar and colpal regions respectively; 10, prep. D351/5 24.9 98 2 Y.1458. 11, prep. D351/4 22.2 95.1, Y . 1459.

Figs 12-17. Phimopollenites augathallaensis (Burger) comb. nov. 12, 13, Scanning electron micrographs X 1,500 of specimen in lateral and polar views, showing sexine modification in colpal areas, South Australia, Haddon Downs No. 5 well 465 ft (141.4m), prep. D351/4 22.1 97.2, Y.1460. 14, Scanning electron micrograph X 4,000 showing sexine sculpture and free-standing columellae in colpal region, Bathurst Island, Meadinga, sample Ni 102, prep. S340/13 21 6 95 1 h i A i l -A 5 ; n ^ f x vi ew Bathurst Island, Moonkinu, sample Ni 130, prep! S 3 4 f / 2 ? } ' 2 J 2 3 > Y - 1 4 6 2 - 16> p olar view of tetracolpoidate specimen, loc. 2 9 9 Xh p r e p ' D 3 5 9 / 1 TSI fifS }}' 94.1, Y.1463. 17, Lateral view, Bathurst Island, Moonkinu, sample Ni 135, prep. S345/1 29.2 87.3, Y.1464. PLATE

4

All figures X 1,000 unless otherwise specified; from unretouched negatives. Figs 1-4.

Figs 5-12.

Tricolpites minutus (Brenner) comb. nov. 1, 2, Polar view, high and median foci, Bathurst Island, Moonkinu, sample Ni 139, prep. S255/1 13 0 100 6 T o i 4 6 5 , l a t e r a l view, high focus, Melville Island, Ant Cliff, sample Ni 280 prep. S330/3 30.9 88.8, Y.1466. 4, Lateral view, high focus, Bathurst Island Meadinga, sample Ni 102, prep. S340/2 28.2 101.9, Y.1467. Tricolpites sp. 5, Lateral view, high focus, Bathurst Island, Moonkinu, sample l V ' V ? r e p ; - S 2 6 4 / 2 2 8 - 4 8 6 - 2 ' Y.1468. 6, Polar view, median focus, Bathurst Island, Moonkinu, sample Ni 139, prep. S255/2 13.0 92.0, Y.1469. 7, 8 Polar view, high and median foci, Bathurst Island, Pulliamandera, sample Ni 185 prep. S335/1 21.9 91.4, Y.1470. 9-12, Scanning electron micrographs of specimen from South Australia, Haddon Downs No. 5 bore, 465 ft (141 4m) nren D351/4 21.9 97.8, Y.1471; 9-11, lateral and polar views X 2,500; 12, detail of sexine in colpal region, X 5,000.

Spec.Publs geol.Soc.Ausf., 4: pp. 3-34, Pis 1-6, 1973.

34

MARY E. DETTMANN Figs 13-19. Tricolpites cooksonae sp. nov. 13-15, Holotype, polar view, high median, and low foci. 16, Polar view, high focus, Bathurst Island, sample Ni 122, prep. S286/2 28.3 91.2, Y.1473. 17-19, Scanning electron micrographs of specimen from Bathurst Island, Patingumputti, sample Ni 193, prep. S301/4 20 1 95.1, Y.1474; 17,18, polar and lateral views X 2,500; 19, detail of sexme X 5,000.

PLATE 5

All figures X 1,000 unless otherwise specified; from unretouched negatives. Figs 1-8. Nyssapollenites lanosus sp. nov. 1, 2, Holotype, polar view, high and median foci 3, 4, Polar view, high and median foci, Bathurst Island, Patingumputti, sample Ni 198, prep. S337/1 22.8 107.5, Y.1476. 5, Lateral view, showing colporoidate apertures, Bathurst Island, Patingumputti, sample Ni 193, prep. S301/2 25.1 101.5, Y.1477. 6, 7, Scanning electron micrographs of specimen X 3,000 in polar and off-lateral aspects, loc. as fig. 5, prep. S301/4 22.2 95.9, Y.1478. 8. Scanning electron micrograph of tetrad X 2,000, loc. as fig. 5, prep. S301/4 22.2 96.1, Y.1479. Figs 9-15. Nyssapollenites squamosus sp. nov. 9, 10, Holotype, polar view, high and median foci. 11, Polar view, median focus, South Australia, Geltwood Beach No. 1 well, core 8, 3,771-91 ft (l,146-52m), prep. K394b/2 20.4 94.1, Y.1481. 12 13, Polar view, high and median foci, Bathurst Island, Pulliamandera, sample Ni 185, prep. S335/1 32.3 96.8, Y. 1482. 14, 15, Scanning electron micrographs of specimen X 3,000 in polar and lateral aspects, Bathurst Island, Patingumputti, sample Ni 193, prep. S301/4 20.1 95.1, Y.1483. Figs. 16-21. Triorites punctulatus sp. nov. 16, 17, Holotype, polar view, high and median foci 18, 19, Lateral view, high and median foci, Bathurst Island, Patingumputti, sample Ni 193, prep. S301/2 16.8 100.3, Y.1485. 20, 21, Scanning electron micrographs of specimen X 3,000 situated in polar and lateral aspects, loc. as figs 18, 19, prep. S301/4 20.6 95.9, Y.1486.

PLATE 6

All figures X 1,000 unless otherwise specified; from unretouched negatives. Figs 1-3 Triorites minor Couper, 1953. 1, Scanning electron micrograph of specimen X 3,000 situated in polar aspect, Victoria, Port Campbell No. 2 well, core 15, 8,407-18 ft (2,555-8m), prep. F079b/5 22.1 96.0, Y.1487. 2, 3. Polar view, high and low foci, Victoria, Port Campbell No. 2 well, core 7, 7,913-30 ft (2,411-7m), prep. F082/3 24.9 113.8, Y.1488. Fies 4-7 Triorites sp. 4, 5, Off-polar aspect, high and median foci, Bathurst Island, ' Pulliamandera, sample Ni 185, prep. S335/3 14.3 91.9, Y.1489. 6, 7, Lateral view high and median foci, Bathurst Island, Patingumputti, sample Ni 193, prep.' S338/1 16.2 92.2, Y.1490. Figs 8-11. Senectotetradites fistulosus sp. nov. 8, Holotype. 9, Single grain in lateral aspect, Bathurst Island, Meadinga, sample Ni 106, prep. S283/2 16.2 104.0, Y 1492 10 11, Scanning electron micrographs of tetrad from Queensland, Mornington' Island No. 1 well, cuttings at 310-20 ft (94-7m), prep. K340/4 21.4 96.1, Y.1493; 11, X 5,000 showing detail of sexine in colpal regions of two grains within the tetrad. Fies 12-16. Senectotetradites varireticulatus sp. nov. 12, Tetrad, X 500, South Australia. Geltwood Beach No. 1 well, core 8, 3,771-91 ft (l,146-52m), prep. F418/2 19 0 84.9, Y.1494. 13, Holotype X 500. 14, Damaged tetrad consisting of two whole and one half grains X 500, Bathurst Island, sample Ni 123, prep. S252/2 20.2 86.9, Y.1496. 15, 16, Scanning electron micrographs of specimen from Queensland, Mornington Island No. 1 well, cuttings at 310-20 ft (94-7m), prep. K340/5 21.1 91.2, Y.1497; 16, X 4,300 showing columellate processes supporting surface reticulum.

MARY

E.

DETTMANN

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE 1

PLATE

2

MARY E .

DETTMANN

MARY

E.

DETTMANN

Spec.Pubis geoI.Soc.Aust., 4, 1973.

PLATE 3

PLATE

4

MARY E .

DETTMANN

MARY E .

DETTMANN

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE

5

PLATE

6

MARY E.

DETTMANN

UPPER TERTIARY PALYNOLOGY IN SOUTHERN NEW SOUTH WALES By H E L E N E A. M A R T I N

(With 10 Text-Figures, 7 Tables and 2 Plates) ABSTRACT

Pollen assemblages recovered from bore samples of the Lachlan Formation and its equivalents, which are found in the Lachlan and Murrumbidgee River Valleys of New South Wales, have been described. They have been divided into three phases, based on the dominance of the Myrtaceae, Nothofagus or gymnosperm taxa in the pollen spectra. In the Lachlan River Valley between Cowra and Forbes, the pollen spectra show a complete cycle of change, starting with the Myrtaceae-Casuarina phase (the oldest), followed by Nothofagus-Cyathea, then gymnosperms and finally ending with a second Myrtaceae-Casuarina phase (the youngest). This pattern of change can be traced in the deposits at Wagga, but the oldest MyrtacGaQ-Casuarina phase is missing. The cycle also exists in the upper levels at Narrandera but it lacks the Nothofagus phase. Comparable assemblages at Hay are poorly represented and not typical. The deeper levels at Narrandera and Hay contain abundant Nothofagus of the brassi pollen type. These assemblages differ markedly from the Nothofagus phase which has only the menziesii and fusca pollen types. Clearly, the deeper levels are older and are tentatively regarded as Miocene in age. The upper levels containing the cyclic change of the three phases, are tentatively regarded as Pliocene in age. Stratigraphically, the Nothofagus and gymnosperm phases are very important as their vertical range in the section is limited. The last appearance of the brassi pollen type of Nothofagus is well defined and is also valuable for correlation. Some of the pollen types in these assemblages are identified with living taxa for which the climatic tolerances are known. These are used, together with the available independent information, to reconstruct the changes in the vegetation and the climate. Migration patterns which would account for these changes have been postulated. INTRODUCTION When this study was started, there was one published work on eastern Australian Upper Tertiary pollen floras. Cookson (1954), in her paper The Cainozoic occurrence of Acacia in Australia', described assemblages from several localities in Victoria. In these she recorded three gymnosperm taxa, six dicotyledonous pollen types, two ferns, an alga, and fungal hyphae and fruiting bodies. Harris (1971) has since described Pliocene assemblages from the Hamilton and Cavendish districts of Victoria. Small collections of macroscopic plant remains found in Victoria had been described previously by Deane (1904) and Patton (1928). At best, these works are a very meagre account of Upper Tertiary floras. The material used in this study has been obtained from bores sunk by the Water Conservation and Irrigation Commission of New South Wales, during a programme of explora-

tion for ground water. Although the Lachlan River Valley between Cowra and Forbes, and the Wagga district on the Murrumbidgee River have been explored most extensively, the programme has also been extended to Narrandera and Hay (see Locality Map, Fig. 1). While this drilling programme has been a valuable source of material, there are several features of these deposits which are usually considered as undesirable for palynological studies. Most of the section consists of sands and gravels which are completely barren. The pollen-bearing carbonaceous clays occur in small lenses and beds of very limited size. The sections obtained are thus discontinuous. Although these carbonaceous clay lenses show evidence of paludal deposition, the influence of river action cannot be entirely discounted, and large parts of the section are of fluviatile origin. These sediments do not contain any direct evidence which can be used to date them. The ages tentatively assigned to them

Spec.PubIs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

36

HELENE A. MARTIN

have been determined from indirect evidence, correlation and the geology of the whole of the western slopes and plains. In spite of these disadvantages, the floral assemblages present certain patterns which can be recognised throughout a whole district, and are repeated from one river valley to another. This study has been undertaken with the view that the available material can be worked, using suitable methods, and interpreted to the best of the available background information, taking into account the limitations and shortcomings. It is hoped that future evidence will clarify much of the tentative nature of this study. GEOLOGY The unconsolidated sediments of the Lachlan River Valley in the Cowra-Forbes district have been divided into two units which Williamson (1964) has provisionally named the Cowra and Lachlan Formations. The Lachlan Formation reaches a maximum thickness of about 90 m and the overlying Cowra Formation, resting on an erosion surface reaches about 30 m. Both consist of interlensed and interbedded gravels, sands, silts, and clays,

but whereas the gravels and sands of the Lachlan Formation are almost entirely quartzose, the Cowra Formation contains the various resistant rock types found today in the catchment area. This important difference indicates that the two formations were derived from different sources; the Lachlan Formation from a quartz-rich source, probably extensive gravel beds which are only found today as remnant hill-cappings (Glen Logan Gravels), whereas the Cowra Formation was derived from direct erosion of the rocks in the catchment area. From the geology of the western slopes as a whole, Williamson (op. cit.) considered that the Cowra Formation is Pleistocene and the Lachlan Formation Pliocene. Equivalents of the Lachlan and Cowra Formations can be distinguished at Wagga where the total thickness of the unconsolidated sediments is a little less, about 80 m. They can also be distinguished at Narrandera, but whereas the Cowra Formation is about the same thickness, sediments similar to the Lachlan Formation are much thicker (as much as 160 m). The pollen floras (discussed later in this paper) show that the deeper parts

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W. 37 at Narrandera are conspicuously older than SAMPLES AND METHODS either Wagga or the Lachlan River localities. from eight bores have been selected At Hay, the unconsolidated sediments are forSamples but material from over 30 even thicker, extending to a depth of about boresthishas study, been All but one have 400 m, but they do not show a clear division been drilled by examined. the percussion or cable tool into two formations. The stratigraphic relation- method. Casing either follows closely behind ships between the Lachlan and Cowra Forma- or is driven ahead of the drilling bit. Water tions and their equivalents in the Murrum- may be added to make a slurry, drilling bidgee Valley, and with the time-equivalent mud is not used. Slurry samples arebutrecovered and older units of the Murray Basin {e.g., from just ahead of the casing or from within at Hay) have not yet been defined. StratiHay bore has been drilled by a rotary graphic analysis of recent drilling information it. Theand samples taken from cuttings. is expected to be published shortly (D. plant Cores from percussion bores provide the Woolley, pers. comm.). reliable subsurface samples. In the Pels (1960) has described the geology of most absence of cores, slurry samples are of conthe Murrumbidgee Irrigation Area and sur- siderable value as the casing prevents contamirounding districts which are immediately north- nation by cavings from strata higher in the west of Narrandera. There are two distinct Cuttings obtained during rotary drillfacies of deposition, lacustrine being overlain section. ing are less satisfactory as they may be conby fluviatile deposits. It appears that extensive taminated with both drilling mud and cavings. faulting and a gradual sagging movement along Only upper limit of the range of a species the fault planes created a number of basins can betheobtained reliably from cuttings. or fault depressions in which lacustrine sediAll of the samples examined are unconsoliments accumulated. Subsequently, fluviatile de- dated silts with a variable content of sand, position has buried this landscape. It is uncer- clay and carbonaceous Treatment tain when movement occurred, but one of follows the acid digestion material. method using HC1, the western faults is part of a major linea- HF, HNO and K C0 . Special care has been ment of the Murray Basin and in this region, to prevent over-oxidation by HN0 , some of the lacustrine deposits have been dis- taken and the subsequent treatment with K C0 placed also. To the east, the lacustrine surface has been the minimum required to clear appears to have remained undisturbed and material. The residues have been mounted the there are several leucite-basalt outcrops resting glycerine jelly containing a little safranin inO on these sediments. The basalts in these dis- dye and the cover slips (No. 0) sealed with tricts are believed to be of Late Pliocene age. nail varnish. On this basis, the lacustrine sediments are re- clear Approximately hundred spore and garded as Tertiary and the fluviatile as pollen types have one been recovered and these Quaternary (Pels, op. cit.). It is not known are described elsewhere (Martin, in press). how far the system of fault depressions exnames of new species not yet published tended beyond the Murrumbidgee Irrigation The cannot be used in this paper, hence these taxa Area or how much of the Upper Tertiary was are referred to by code numbers. A combinadeposited under lacustrine conditions. This tion of qualitative and quantitative methods point is important for palynology. Fluviatile has been adopted as the best procedure for deposits have a poor reputation as a source analysis of these assemblages. material as the pollen in the sediments could have originated anywhere within the catch- THE POLLEN SPECTRA ment area. Lacustrine sediments are much preferred so that the possibility of long-distance The Lachlan River Valley In the Cowra-Forbes district, the Lachlan transport by water is eliminated. Although Formation extends from below Forbes, upthese deposits have been formed in conjunction stream to Cowra and along Back Creek for with river systems, they are not entirely nearly 40 km. Figure 2 shows the drilling fluviatile. The carbonaceous clay lenses, often sections of the exploration programme undercontaining iron pyrites nodules and dissemina- taken by the Water Conservation and Irrigations, are evidence of paludal deposition. It tion Commission of New South is these carbonaceous clays that are particu- Material from Sections 3 to 8 has Wales. been larly suitable for pollen analysis and have studied. Section 6 has yielded the maximum been used in this study. amount of information about the pollen spectra Spec.Publs geol.Soc.Aust, 4: pp. 35-54, Pis 1-2, 1973. s

2

3

3

2

3

HELENE A. MARTIN 38 (shown in Fig. 3). The depth is indicated to botanical affinities and these groupings are and where several consecutive samples have listed in Table I. From Figure 3, it can be seen that the yielded similar spectra, the average is plotted and this is indicated by the brackets. The oldest and deepest samples have high proporspores and pollen have been grouped according tions of Myrtaceae and Casuarina. Compositae

FORBE

<$>C

Drilling Section

wmi 8

16 km

Oct> L]

:OWRA

j

Scale

Fig. 2. The drilling cross sections of the Lachlan River Valley. Each section has 10-15 bores sunk to basement rock. Data provided by courtesy of the Water Conservation and Irrigation Commission of N.S.W. I

DEPTH (m)

48 65 67

TOTAL SPORES

GYM

CAS

MYRT

Scale :

0 I

40

NOTH m+f

PHASE MYRT - C A S GYM NOTH - C Y MYRT - C A S

COMP GRAM

80 %

l 1 1 I of total pollen count

Fig. 3. The pollen spectra of the Lachlan River Valley, from a bore in Section 6. Where several consecutive samples have yielded similar spectra the average is plotted and this is indicated by the brackets. Abbreviations: GYM, gymnosperms; MYRT, Myrtaceae; CAS, Casuarina; NOTH, Nothofagus; m, the menziesii pollen type; f, the fusca pollen type; COMP, Compositae; GRAM, Gramineae; CY, Cyathea; P, Podocarpaceae; A, Araucariaceae. See Table I for explanation of groupings. For explanation of the pollen spectra, see text.

U P P E R T E R T I A R Y P A L Y N O L O G Y IN S O U T H E R N N.S.W. TABLE I

The groups of spores and pollens

Group Total spores

Natural affinities Lycopodium sp. Osmundaceae Gleichenia sp. Cyathea spp. Hypolepis sp. Phyllocladus Dacrydium sp. Podocarpus sp. Podo carpus sect. Daerycarpus Microcachrys sp.

Phyllocladidites palaeogenicus Cookson & Pike 1954a. Dacrydiumites florinii Cookson & Pike, 1953Z>. Podocarpidites ellipticus Cookson, 1947. Dacrycarpites australiensis Cookson & Pike, 1953a. Microcachryidites antarcticus Cookson, 1947. Araucariacites australis

Araucariaceae

Myrtaceae Casuarina

Nothofagus

Cookson, 1947. All species of Myrtaceidites.

All taxa within the family Casuarina spp.

Casuarinidites cainozoicus Cookson & Pike, 1954b. Triorites harrisii Couper, 1953. Nothofagidites aspera Cookson, 1959. Nothofagidites brachyspinulosa

Nothofagus menziesii pollen type.

Nothofagus fusca pollen type.

Compositae

All taxa within the family.

Gramineae

All taxa within the family.

and Gramineae are at their maximum for the sequence but the frequencies are low in comparison with the other groups. At 78 m the total spores reach high frequencies. Cyathea spp. (CY) account for about half, as indicated by the cross line. Coinciding with the total spore peak, Nothofagus reaches its highest frequency and most of the pollen is the fusca type (f). Myrtaceae and Casuarina have both reached their lowest frequencies and Compositae and Gramineae are not present at this level. Above this, the frequency of Nothofagus declines rapidly, the total spores more gradually, whereas Myrtaceae and Casuarina increase. At this level, the gymnosperms reach their highest frequency. Most of the pollen belongs to the Podocarpaceae (P) with only a small portion of Araucariaceae (A). Finally, in the uppermost (youngest) sample, Myrtaceae and Casuarina are again the dominant groups.

Palynological classification

All forms of spores, mainly pteridophytes but including some bryophytes.

Cupressaceae

Gymnosperms

39

Cookson, 1959.

Tubulifloridites antipodica Cookson, 1947. Tubulifloridites sp. (spp. ?) Graminidites media

Cookson, 1947.

The spectra have been divided into three phases based on the dominant groups and this is also shown in Figure 3. The MyrtaceaeCasuarina phase occurs twice, as the oldest and youngest phases. Associated with these phases is a number of pollen types found in low frequencies (3 per cent or less). These are listed in Table II and described in the Appendix. Trilites bifurcatus Couper, 1960 (probably a hepatic), Acacia myriosporites Cookson, 1954, Microntheum sp., Forms Q-32, and R-28 are found in the Myrtaceae-Cayuarina phase, both older and younger. Drimys sp., Quintinia sp., and Form P-30 are associated with the Nothofagus phase. The gymnosperm phase has Drimys sp., Form R-28; less frequently, Trilites bifurcatus, Micrantheum, Form Q-32 and, rarely, Acacia. In this respect, the gymnosperm phase is intermediate between the Nothofagus and

Spec.Publs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

HELENE A. MARTIN

40 TABLE I I

in Sections 3, 6, and 8, and Figure 4 shows the spectra f r o m these. It is located at greater depths, downstream, as total depth of the Lachlan Formation also increases. The variation between the spectra from each section is small, but there is a steady decline of the Nothofagus content, f r o m upstream to downstream. The gymnosperm phase has been located in Sections 4, 5, and 6, and shows similar patterns in that the frequencies also decrease, downstream. Whereas there are only two taxa in the Nothofagus group, and the proportions are shown in Figure 4, there are seven in the gymnosperms. The frequencies of these are shown in Table III. The presence of Phyllocladus in such high frequencies in Section 5 is surprising as it is almost entirely absent from all other samples of the Lachlan Formation.

Lachlan River Valley: the low frequency taxa (3 % or less of total spore-pollen count)* Depth of Sample, Phase (see Fig. 3) (m) 48

Taxon

MyrtaceaeCasuarina (younger)

Trilites bifurcatus Acacia myriosporites Micrantheum Form Q-32 Form R-28

Gymnosperms

Trilites bifurcatus (67 m only) Drimys Micrantheum (67 m only) Form R-28 Form Q-32 (67 m only)

65 67

78

Nothofagus-Cyathea Drimys Quintinia Form P-30 Form B-17

79

MyrtaceaeCasuarina

Comparison with the Cowra Formation Some pollen spectra f r o m the overlying Cowra Formation have been described (Martin, 1969). When compared with those of the Lachlan Formation, they have a much greater quantity of Compositae pollen, including two

Trilites bifurcatus

82 85

Acacia myriosporites Micrantheum Form Q-32 Form R-28

TABLE I I I

* For descriptions of code-numbered taxa, see Appendix.

Lachlan River Valley: frequency of the gymnosperm taxa (% of total spore-pollen count) Section (See Fig. 2)

MyrtaceaQ-Casuarina phases. Haloragis (Haloragacidites haloragoides Cookson & Pike, TAXON (See Table 1) 1954b) is present in each phase and does not Cupressaceae show any particular association. Phyllocladus The My TiSiCQSLQ-Casuarina phase is veryDacrydium Podocarpus common and occurs in every section shown Podocarpus sect. Dacrycarpus in Figure 2 and every bore examined. Samples Microcachrys containing the Nothofagus and gymnosperm Araucariaceae phases are few, and are restricted to certain TOTAL depths. The Nothofagus phase has been found DEPTH

4

5

2

—

44

26

21-5

SECTION

(m)

77-79

98-99 MYRT

GYM SCALE :

0

1

CAS

20

1

40

NOTH

m• f

COMP

—

5 14-5 12-5 1 9

— 2 10 7 1 2-5

61-64

TOTAL SPORES

6

8 6-5 6-5 4-5 — 0-5

—

GRAM

60 % of total pollen count

1 1

Fig. 4. The Nothofagus Phase from Sections 3, 6 and 8 (see Fig. 2). The abbreviations used are the same as for Figure 3. For explanation see text.

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W. DEPTH

41

SECTION

(m) 57-60

i

65-67

64-67 TOTAL SPORES

GYM

MYRT

20

SCALE

0

CAS

COMP

GRAM

SO % of total count

Fig. 5. The Gymnosperm Phase from Sections 4, 5 and 6 (see Fig. 2). The abbreviations used are the same as for Figure 3. For explanation, see text. additional kinds not found in the latter, and the frequency may exceed 60% of the total count. Gramineae pollen is also more abundant, 20% or more. Of the other groups of spores and pollen, the total spore count is low, the gymnosperms rare or absent, Myrtaceae common, Casuarina low, and Nothofagus absent. Thus, the spectra of the Lachlan Formation are quite distinct from those of the Cowra Formation. Wagga The unconsolidated sediments in the valley at Wagga are considered equivalent to the Cowra and Lachlan Formations. The resistant rock type of the latter at Wagga is almost entirely quartzose, as in the Lachlan River Valley. The pollen spectra from the bore with the best sequence are shown in Figure 6. They are similar to those of the Lachlan River DEPTH

(m) 37

Valley, with relatively minor variations which are discussed below. The total spores show two exceptionally high counts of over 50% and 80% at depths of 39 m and 64 m respectively. However, other bores, including one only 100 m away, did not yield excessively high spore counts, which suggests that these result from localised deposition and are not representative of the regional spore-pollen flora. Such a high fern spore frequency distorts the rest of the spectrum and the quantitative aspects of these samples are best disregarded. The taxa which make up the spore group are more diversified than those of the Lachlan River Valley. Rare forms in the latter are more common at Wagga, e.g. Form B-17 (see Appendix), and additional kinds not seen at the former location include Cyatheacidites annulata Cookson, 1947. PHASE

^

MYRT - CAS

m ,t

5T

| GYM J NOTH - C Y

GRAM ICAS m +' COMP 40 80 % of total count Fig. 6. The pollen spectra at Wagga, from a bore about 2 km E.S.E. of Wagga Wagga township. The abbreviations used are the same as for Figure 3. For explanation, see text. Spec.Publs geol.Soc.Aust, 4: pp. 35-54, Pis 1-2, 1973. GYM

SCALE :

MYRT

, w ' ; '

HELENE A. MARTIN TABLE V The gymnosperm frequencies show a welltaxa {less than 3% of total defined peak. The counts of Myrtaceae are Wagga: low frequency spore-pollen population)* generally lower than those of the Lachlan River Valley, Casuarina is low throughout, and Depth of the Nothofagus peak is sustained through four sample Phase Low frequency taxon (m) samples. Whereas most of the Nothofagus pollen in the Lachlan River Valley is the fusca 27 Trilites bifurcatus type, at Wagga, the menziesii type forms the Acacia myriosporites major portion. Both Compositae and GramiR-28 neae are present and more abundant in the 39 Myrtaceae Cyatheacidites annulata upper part, although the frequencies are still Acacia myriosporites relatively low. Micrantheum The spectra at Wagga can be divided into Drimys 59 the same three phases, as seen in the Lachlan River Valley. The cycle of change follows the Cyatheacidites annulata Gymnosame sequence, except that the lower and older Quintinia 61 sperm Q-32 MyrtacQae-Casuarina phase is missing. The deepest pollen-bearing deposits contain the Cyatheacidites annulata 62 Nothofagus-Cyathea phase which is followed B-17 by that of the gymnosperms. Between the two, Drimys ? Quintinia the 62 m sample has a rather unusual compoQ-32 sition. The count of Myrtaceae is higher than either that of Nothofagus or gymnosperms and Cyatheacidites annulata B-17 Form Q-32, which is usually associated with 64, 66 Nothofagus Drimys the MyrtaceaQ-Casuarina phase, is also present. Quintinia (66 m only) The gymnosperm phase has quite appreciable quantities of Nothofagus and in this respect it * For descriptions of code-numbered taxa, see differs from that of the Lachlan River Valley. Appendix. Table IV presents the frequency of the gymnosperm taxa at Wagga, and they show the same and in another bore at Wagga, Drimys to trends seen in the Lachlan River Valley. 9%. Haloragis occurs down to a depth of The younger Myrtaceae-Casuarina phase is 61 m, but not below it. As already mentioned, present, but with less Myrtaceae, less Casua- some rare species of the Lachlan River spectra rina and more spores than in the Lachlan are more common, and there are additional River spectra. The low frequency taxa from low frequency species. each sample are listed in Table V, and they The Nothofagus and gymnosperm phases show the same patterns as those in the Lachlan can be located in several bores nearby, beRiver Valley. Some occur in surprisingly high tween the 59 m and 64 m levels. The sedifrequencies, e.g. Acacia 11%, Quintinia 4.5%, ments below these depths extend down to 80 m, are sands and gravels, and do not contain TABLE I V These strata at Wagga correlate with Wagga: frequency of gymnosperm taxa (% of total pollen. those of the Lachlan River Valley, between spore-pollen population) the depths of 60 m and 90 m, in which the Depth (m) of sample* Nothofagus and gymnosperm phases are Taxon found. 27 59 61 62 66 Narrandera 1 1 1 Cupressaceae A bore at Narrandera extends down more 0-5 0-5 than 1 Phyllocladus 160 m and penetrates the equivalents of 0-5 0-5 1-5 0-5 Dacrydium the Cowra and Lachlan Formations. For the Podocarpus (P. 1 19 27-5 2-5 4 ellipticus) entire depth below 27 m, the limit of the Podocarpus sect. Cowra Formation equivalent, the resistant rock 1-5 0-5 1 18 Dacrycarpus type is almost entirely quartzose. The pollen 1-5 0-5 2 Microcachrys spectra of these deposits are presented in 1-5 2-5 6 Araucariaceae Figure 7. 9 14 1-5 37-5 35 TOTAL Below 99 m, the spectra are very different to those above. Nothofagus dominates the * The high spore samples have been omitted.

42

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W.

lower part of the profile whereas it is completely absent from the upper part. Most of it is the brassi pollen type (predominantly Nothofagidites emarcida Cookson, 1959), whereas the menziesii and fusca types make up only a small portion. Spores, gymnosperms, and Casuarina are relatively low; Compositae and Gramineae are absent. Drimys and Quintinia are usually present in each sample. There are relatively minor fluctuations up to the 99 m level where the Nothofagus content declines markedly. These spectra, with very high quantities of the brassi pollen type, are characteristic of much of the Lower Tertiary. The brassi pollen type disappeared from the Gippsland Basin of southeastern Australia about the end of the Miocene (Partridge, 1971). However, the climate of these inland districts would not have been comparable to that of the coastal Gippsland Basin, which was probably the most favourable for continued existence of Nothofagus. Consequently, the disappearance of the brassi pollen type in these deposits at Narrandera could have occurred before the end of the Miocene. At the 99 m level, Myrtaceae becomes the dominant pollen group. About half of it is Myrtaceidites parvus Cookson & Pike, 1954b, and there is also some Myrtaceidites mesonesus Cookson & Pike, 19546. Myrtaceidites eucalyptoides Cookson & Pike, 19546, the eucalypt pollen type, forms about one tenth of the myrtaceous count. It also occurs below this

level, but it is rare. Acacia first appears at this depth. Micrantheum sp., Haloragis, and Form Q-32, commonly associated with the Pliocene spectra of the Lachlan River and Wagga, are not found at the 99 m depth or below it. Form R-28 is present at 99 m. Above the 99 m depth, the spectra are typically those of the MyrtaceaQ-Casuarina phase of the Lachlan River Valley. Myrtaceae is by far the most abundant group, and Casuarina is low, but shows an increase over that below it. Nothofagus is absent, gymnosperms are at their lowest, and both Compositae and Gramineae are present. Micrantheum, Haloragis, and Form Q-32 are first noted at 54 m in this bore. Unfortunately, the section between 54 m and 99 m does not contain pollen-bearing material, and this is a substantial gap. However, four samples at depths of 70, 72, 73, and 74 m, from another bore 3.6 km to the south, have all yielded the Myrtaceae-Casuarina phase, and the low frequency taxa listed above are also present. Although the 70-74 m level of this second bore is considered as intermediate to 99 m and 54 m in the first bore, there is always some doubt about the correlation of depths in these sediments. There is a distinct gymnosperm peak at 34 m and 36 m, but Nothofagus does not reappear at these levels or immediately below, as we would expect from the spectra of the Lachlan River Valley and Wagga. The gymnosperm frequency, down to 99 m, is presented

DEPTH

PHASE (of Pliocene) MYRT-CAS

(m) 32 3 4 <* 36 40

GYM MYRT-CAS

54 99 115 <1 12 4 <1 126

132 142

MIOCENE

149 150 <1 161 <1

TOTAL SPORES

NOTH (m+f) + b

GYM

s c a l E

•

0

^

40

COMP

GRAM

80 % of total count

Fig. 7. The pollen spectra at Narrandera, from a bore about 1£ km S.S.W. of Narrandera township. The abbreviations used are the same as for Figure 3. For explanation, see text. Spec.Publs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

HELENE A. MARTIN

44

in Table VI, which shows that two of the gymnosperm taxa (Phyllocladus and Dacrydium) have not been recorded from this part of the section. Podocarpus (of the P. ellipticus type) and the Araucariaceae are the only two taxa present in any abundance. Casuarina, Compositae, and Gramineae show a steady increase towards the top of the sequence, which ends with the younger MyrtacQae-Casuarina phase with Casuarina the dominant. Both Compositae and Gramineae show a far greater TABLE V I

Narrandera: frequency of gymnosperm taxa (% of total spore-pollen population) Depth of sample (m) Taxon 32 34 36 40 54 99 Cupressaceae 0-5 2 Podocarpus (P. ellipticus) 2- 5 21 22 7 0-5 3 Podocarpus sect. Dacrycarpus 0-5 0-5 Microcachrys 1 0-5 Araucariaceae 1 7-5 5-5 1*5 2-5 23 29-5 13-5 1 7 TOTAL

increase towards the top of the sequence when compared with the previous two localities. The upper part of the sequence contains some low-frequency species not found elsewhere. These include Tricolporopollenites pelargonioides sp. nov., Tricolpites geranioides Couper, 1960, and the Portulacaceae (see Appendix) . Cyatheacidites annulata and Form B-17, which are found at Wagga, do not occur in the Narrandera sequence, and the fern spore flora is quite restricted. The low-frequency taxa are listed in Table VII. Hay

A bore at Hay extends to a depth of almost 210 m through deposits in which the equivalents of the Cowra and Lachlan Formations cannot be identified. Almost the entire section consists of clays, which contrasts with the quartzose sand and gravel beds of the other localities. The pollen spectra are presented in Figure 8. The spectra below 115m correlate well with those below 99 m at Narrandera. Nothofagus is again the dominant group and the brassi pollen type forms the largest portion. The

TABLE V I I

Narrandera: low frequency taxa (less than 3 % of total spore-pollen population) Taxon Depth of Sample Phase and epoch* (m) Trilites bifurcatus Myrtaceae-Gzswtfrwtf phase, 32 Drimys Micrantheum Tricolporopollenites pelargonioides Pliocene Tricolpites geranioides Trilites bifurcatus Gymnosperm phase, 34, 36 Drimys Micrantheum Pliocene T. pelargonioides (36 m only) Form R-28 (36 m only) Trilites bifurcatus (54 m only) Myrtaceae- Casuarina phase, 40, 54 Acacia Drimys (54 m only) Pliocene Micrantheum T. pelargonioides Portulacaceae (40 m only) Form Q-32 (54 m only) Form R-28 (54 m only) Drimys Miocene 99 Quintinia Acacia Form R-28 Drimys Miocene 115-161 Quintinia Form P-30 * These ages are tentative.

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W.

(m)

45

DEPTH 91

POSTTERTIARY ?

<

109

112

PLIOCENE ?

115 118

121

133

4

MIOCENE

151 169

<b

TOTAL SPORES

MYRT

SCALE:

I

b

(m + f) + b NOTH

CAS 40

gO %

COMP

I

GRAM

CHENOPODTYPE

of total count

Fig. 8. The pollen spectra at Hay, from a bore about 16 km E. of Hay township. The abbreviations used are the same as for Figure 3. For explanation, see text.

upper limit of Nothofagus is uncertain, probably because of recycling from deposits further upstream. Above 115 m, Myrtaceae is the dominant group and Forms Q-32 and R-28 occur at 109 m. This is considered as evidence that the spectra of 109 m and 112 m may be equivalent to the upper part of Narrandera, 54 m and above, although somewhat aberrant. There is a small peak in the gymnosperms (109 m) but it is uncertain whether this correlates with the gymnosperm phase. Compositae first appear at 112 m but Gramineae extends to the deepest sample although it is rare below 112 m. The spectrum at 91 m has a surprisingly high content of chenopod-type pollen (Chenopodiaceae, Amaranthaceae, and Caryophyllaceae). If the chenopod-type content is disregarded, Compositae would dominate the spectrum which could then correlate with those of the Cowra Formation. The chenopod-type occurs in the Cowra Formation, but at much lower frequencies (to 7%). It is unlikely (though not impossible) that the chenopodtype has been carried down during the drilling process, since two samples above 91 m are entirely devoid of pollen. If this spectrum does correlate with the Cowra Formation, then it occurs at a much greater depth. Elsewhere the Cowra Formation does not exceed 30 m. Clearly, the 91 m level does not correlate with the spectra of the other localities. It may be post-Tertiary but further evidence is required to substantiate this suggested age.

Other localities At Dubbo, on the Macquarie River, two samples at depths of 30 and 33 m from the same bore have assemblages most like the gymnosperm phase of the Lachlan Formation. These samples also contain a small amount of Nothofagus of the menziesii pollen type. At Albury on the Murray River, samples from depths of 51 to 60 m have yielded the Myrtaceae-Casuarina phase, and there are some gymnosperms present as well. This has been the only material examined from these rivers and it is likely that further exploration in the Macquarie and Murray River Valleys will show that a comparable sequence is present. Material from 6 bores at Gwyder River localities (Moree, Biniguy) has failed to yield Upper Tertiary assemblages. Over 20 bores from the Namoi River (Wee Waa to Gunnedah), have been examined, and whereas Mesozoic assemblages are frequently recovered, only one of the Upper Tertiary has been found. This assemblage from 81 m at Boggabri, can be correlated with the MyrtaceaQ-Casuarina phase of the Lachlan Formation. CONCLUSIONS Summary of the stratigraphic implications The Nothofagus and gymnosperm phases are extremely important for correlation as their vertical extent in the section is limited. If located, they can be used as marker bands. The last appearance of the brassi type of Nothofagus is also important for correlation. Spec.Publs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

HELENE A. MARTIN The pollen spectra of the Upper Tertiary positae and Gramineae are common also. units are summarised in Figure 9. Whereas Casuarina is found throughout the region the different strata are correlated with confi- (Beadle, 1948). Of the fossil flora, Nothofagus, six of the dence (unless otherwise indicated), it must be stressed that the ages are tentative and await seven gymnosperm taxa, Drimys and Quintinia are not found in these regions today. The positive evidence. The best development of the supposed Upper Tertiary sequence is seen in menziesii pollen type of Nothofagus occurs in disjunct communities in Victoria (N. cunningthe Lachlan River and Wagga spectra, where both the Nothofagus and gymnosperm phases hamii (Hook.) Oerst) and the east coast tableare present. Narrandera shows a modification lands of New South Wales (N. moorei (F. in that the Nothofagus phase is missing, Muell.)) Maiden of the Barrington Tops, (exwhereas at Hay, a comparable sequence is tending to southern Queensland). Its nearest inadequately represented and not typical when occurrence to these localities where the fossils compared with the other locations. The pre- have been found, is about 300 km to the sumed Miocene sequence is only found at northeast or more than 300 km to the south. The fusca pollen type of Nothofagus, and the Hay and Narrandera. gymnosperms Phyllocladus, Dacrydium, Podocarpus sect. Dacrycarpus, and Microcachrys Phytogeographic considerations When comparing these pollen assemblages are not found on the Australian mainland with the flora found in this area today, there today. Podocarpus, the P. ellipticus type, and are some notable discrepancies. In the eastern Araucariaceae grow in coastal districts, and part, eucalypt woodland, with a herbaceous the nearest occurrence of the latter is over ground cover containing many species of 300 km to the northeast. Of the gymnosperm Gramineae and Compositae, is common. Dry taxa, only the Cupressaceae is found in these sclerophyll forest, which has species of euca- localities, but this is discussed further, below. Nothofagus, the gymnosperms, Drimys, and lypts as dominants, a well-developed sclerophyllous shrub layer, and lacks a herbaceous Quintinia are usually found only in rain-forest ground cover, has a restricted distribution. To communities. Today, rain-forest of any kind the west, scrub communities are found, to- does not occur west of the divide (Baur, gether with eucalypt woodland. There are 1957), which is about 100 km east of Cowra. many different species in the scrub and the In the Cupressaceae, there are at least two dominants may be mallee eucalypts, Acacia species of Callitris which grow in eucalypt spp. or chenopodiaceous shrubs, and Com- woodland and dry sclerophyll forest, and are

46

DEPTH

DEPTH

T 0

(m)

NARRANDERA —

WAGGA

F

LACHLAN

RIVER

>

- GYM

HAY

NOTH^ POSTTERTIARY/ • ?

~ MY RT-

_ _

_

GYM MYRT

NOTH

"

. N0TH "

6YM MYRT

NOTH

- MYRT

PLIOCENE

-

_—

—

—

MYRT

NOTH

•

MIOCENE

0

40

8 0 % OF T O T A L

POLLEN

COUNT

MIOCENE

GYM

J

ABBREVIATIONS i

6YM

-GYMNOSPERMS

MYRT

- MYRTACEAE

NOTH

- NOTHOFAGUS

Fig. 9. Summary of the stratigraphic implications. The ages quoted are tentative.

(m>

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W. 47 found in these districts, but Callitris mac- brassi pollen type of Nothofagus and many leayana (F. Muell.) F. Muell. of southern other Lower Tertiary forms do not reappear. Queensland, occurs in rain-forest. The fossil This must have been a response to increased Cupressaceae is found only in association with precipitation, which allowed the rain-forest of the other gymnosperms and Nothofagus, which the refuge areas in the highlands to expand suggests that it was a rain-forest taxon and and migrate down-stream. However, it did not is thus not comparable with the species found entirely replace the Myrtaceae-Casuarina phase in this region today. as these taxa occur with Nothofagus and the Although Eucalyptus spp. are the dominants gymnosperms. Most likely, there was a mosaic, and Casuarina is found throughout this whole similar to the mosaics which are a feature of region, the MyrtaceaQ-Casuarina phase cannot the vegetation today. Distance from the refuge be equated with the vegetation found there areas may have had an effect on these postutoday. The eucalypt pollen type is present in lated migrations, but the difference in the the pollen assemblages, but is not particularly Nothofagus and gymnosperm phases seen at abundant. Myrtaceidites mesonesus Cookson Wagga and Narrandera, may have resulted & Pike, 1954b, is the most abundant fossil also from a climatic gradient, similar to that type, and McWhae (1957) suggests that it seen today. Climatic conditions favourable to bears a close resemblance to Whiteodendron, rain-forest did not last, and Nothofagus and one of the Tristania complex. Myrtaceidites the gymnosperms disappeared, leaving only parvus Cookson & Pike, 1954b, of uncertain the MyrtacQae-Casuarina phase. affinities, is also quite common. The paucity of These spectra show changes between two Compositae and Gramineae in the spectra is main types of vegetation, rain-forest of the another discrepancy, considering the abun- Nothofagus and gymnosperm phases and (predance of these taxa in the region today. sumably) forests of Myrtaceae and Casuarina. If a modern counterpart of these fossil Towards the top of the sequence, there is eviassemblages is sought, part of it could be dence of a third type. The increase in Compofound in New Zealand. The two kinds of sitae and Gramineae at Narrandera probably Nothofagus and all of the gymnosperm taxa indicates the development of the herbaceous grow there today. However, New Zealand does ground cover, and possibly savannah, which not have a counterpart of the Myrtaceae- are features of these regions today. Casuarina phase, which should be sought in Climatic interpretations Australia. The climatic requirements of the taxa identiA tentative outline of the changes since field in these assemblages can be used to postu(presumed) Miocene time can be recon- late changes in the climate. There is also some structed. As the pollen spectra form a dis- independent evidence from the lithologies of jointed sequence, there could be large dis- these deposits. The sequence is not continuous, junctions in the events outlined below, and so the possibility of disjunctions, and the tentathis should be borne in mind. The rain-forest tive nature of these interpretations should be floras, in which the brassi pollen type of borne in mind. Nothofagus is so conspicuous, disappeared In New South Wales, rain-forest has a very from these districts, presumably about Miocene restricted distribution and the requirements for time. No doubt this was a response to de- its existence are well known. The most imporcreasing precipitation, and this is discussed tant factor is a high level of precipitation, and further in the following section. However, at least 150 cm per annum are required for some of the taxa must have survived in refuge widespread development of rain-forest. Where areas, most likely in highlands to the east, the precipitation is strongly seasonal, dry rainwhere precipitation would have been higher. forest results (Baur, 1957). At critical levels, Most of the taxa of the Myrtaceae-Coswarma when precipitation is limiting, rain-forest is phase are present in these older assemblages, confined to sheltered slopes and the protected but in low frequencies. They became abundant gullies which maintain a higher humidity. after the disappearance of the rain-forest. Edaphic factors are also important, and rainDuring presumed Pliocene time, the rain- forest is usually found on the deeper, heavier forest reappeared in these districts, and all soils which have a greater storage capacity for of the taxa associated with the Nothofagus water. These soils are generally more fertile as and gymnosperm phases are found in the well. Lower Tertiary assemblages. However, the Most of the rain-forest in New South Wales SpecJPubls geolJSoc.Aust, 4: pp. 35-54, Pis 1-2, 1973.

HELENE A. MARTIN

48

or slightly above them. There was another sharp drop at what is thought to be the end of the Pliocene and the beginning of the Pleistocene (Dorman & Gill, 1959; Dorman, 1966). The level of precipitation can be deduced from the requirements of the taxa identified in the fossil floras. For the older (presumed Miocene) assemblages containing the brassi pollen type of Nothofagus, precipitation must have been at least 150-180 cm per annum and possibly more. The disappearance of this assemblage was probably associated with a decrease in precipitation to below 150 cm, although a drop in temperatures may have been involved as well, as indicated by the palaeotemperatures. The Nothofagus and gymnosperm phases resulted from an increase in precipitation, to at least 150-180 cm. The greater development of carbonaceous clays, often containing iron pyrite disseminations and nodules, which occur together with these phases, substantiate this interpretation. There

is classified as subtropical, but Nothofagus is found in the montane or temperate rain-forests. The latter are restricted to higher altitudes where temperatures are lower, precipitation is higher (180 cm per annum or more), and mists and fogs are common (Baur, 1957; Frazer & Vickery, 1938). Considering the Nothofagus and gymnosperm forests in New Zealand, precipitation is not a limiting factor but temperature, as determined by altitude, has a marked influence on distribution. Gymnosperms are a feature of the warm temperate belt whereas Nothofagus becomes significant in the cool temperate (or montane) and subalpine belts (Wardle, 1964). Independent evidence of temperature changes can be obtained from oxygen isotope palaeotemperatures, although there are some divergent values and the results are rather meagre. There was a fairly sharp drop from about the Oligocene to Middle Miocene, after which temperatures remained at about the same level, comparable with those of today,

1150 - 1 8 0 1

ESTIMATED MAXIMUM

PRECIPITATION

DURING THE LATE T E R T I A R Y (cm PUI)

45

PRESENT-DAY PRECIPITATION (cm p.a.)

N

A

. 5

FORBES

**1150-I80l 63*COWRA \

*

SCALE

—i80

1

160 km

Fig. 10. Estimated maximum precipitation at the time of the Nothofagus and gymnosperm phases (tentatively Late Pliocene) compared with present-day values. For explanation, see text.

UPPER TERTIARY PALYN< LOGY IN SOUTHERN N.S.W. 49 may have been an initial decrease in tempera- gramme). The highlands to the east have also ture, judging from the relationships of Notho- existed throughout this time, being maintained fagus and the gymnosperms in New Zealand. by episodic uplift (Brown, Campbell & Crook, Precipitation did not remain at this level but 1968). If, during the Late Tertiary, the main decreased once again, as indicated by the return moisture-bearing winds came predominantly of the MyrtaceaQ-Casuarina phase. The pre- from the east, as they do today, then there dominantly grey colour of the whole of the would have been a progressive decrease in sediments (Lachlan Formation and equiva- precipitation westward, once they passed over lents) indicates that there was no marked dry the highlands. A well-marked dry season, which season. is characteristic of these inland areas today, The Nothofagus and gymnosperm phases are has developed subsequent to the time of these well represented in the Lachlan River Valley fossil assemblages. and Wagga. In the latter there is a greater Burbidge (1960), in discussing the phytodiversity of spores, including species not found geography of the Australian region, remarks elsewhere {e.g. Cyatheacidites annulata). Most on the northwards retreat of the rain-forest of the spores are pteridophytes and a well- during the Pliocene. This study shows that developed fern flora usually requires high there have been east-west migrations as well, levels of precipitation. Thus at the time of these and it is postulated that these have been a phases, Wagga may have had a greater pre- response to changes in the level of precipitacipitation than that of the Lachlan River tion. Burbidge also suggests that there was a Valley. On the other hand, Narrandera lacks swing back to subtropical conditions during the Nothofagus phase, the gymnosperms are the Pliocene, but does not indicate the basis somewhat restricted and the spore flora is of this assumption. This study provides eviparticularly limited. This is considered as dence of a period of increased precipitation evidence that the precipitation was not suffi- which is tentatively regarded as Pliocene, and cient for rain-forest and it did not reach 150 possibly Late Pliocene. cm per annum. The estimated precipitation at the time of the Nothofagus and gymnosperm ACKNOWLEDGMENTS phases is presented in Figure 10, together with The Water Conservation and Irrigation those of the present day. Commission of New South Wales has generThe present-day precipitation shows an east- ously supported this project. Messrs. W. H. west gradient, higher to the east and decreasing Williamson and D. Woolley of the Commission westward. The fossil floras show evidence of have given invaluable assistance. I am indebted this gradient also, though at a higher level, to the late Professor H. N. Barber for his enas seen in Figure 10. This is not surprising. couragement of this work and to Professor D. These rivers have always flowed westwards, Walker, for his invaluable advice. I am gratealong approximately the same course as today ful to the staff of the School of Botany who (evidence from the exploratory drilling pro- have assisted in many ways. REFERENCES BAUR, G. N., 1957: Nature and distribution of

rainforest in New South Wales. Aust. J. Bot.,

Plant microfossils from the Kerguelen Archipelago. Rep. B.A.N.Z. Antarct. Res. Exped., A2, pp. 127-

COOKSON, ISABEL C., 1947:

5, pp. 190-233. BEADLE, N. C. W., 1948: The vegetation and pas-

tures of western New South Wales. Dept Conservation, N.S.W.

BROWN, D . A., CAMPBELL, K. S. W., & CROOK,

K. A. W., 1968: The Geological Evolution of Australia and New Zealand. Permagon Press, Oxford. BURBIDGE, NANCY T., 1960: The phytogeography of the Australian region. Aust. J. Bot., 8, pp. 75-212.

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142.

, 1954: The Cainozoic occurrence of Acacia in Australia. Aust. J. Bot., 2, pp. 5259. , 1957: On some Australian Tertiary spores and pollen grains that extend the geological and geographical distribution of living genera. Proc. R. Soc. Vict., 69, pp. 41-54. , 1959: Fossil pollen grains of Nothofagus from Australia. Proc. R. Soc. Vict., 71, pp. 25-30.

50

HELENE A. MARTIN

, & PIKE, KATHLEEN M . , 1953a: The Ter-

tiary occurrence and distribution of Podocarpus (Section D aery carpus) in Australia and Tasmania. Aust. J. Bot., 1, pp. 71-82. , , 19536: A contribution to the Tertiary occurrence of the genus Dacrydium in the Australian region. Aust. J. Bot., 1, pp. 474-484.

, , 1954a: The fossil occurrence of Phyllocladus and two other podocarpaceous types in Australia. Aust. J. Bot., 2, pp. 6068.

, , 1954b: Some dicotyledonous pollen types from Cainozoic deposits in the Australian region. Aust. J. Bot., 2, pp. 197219. COUPER, R. A., 1953: Upper Mesozoic and Cainozoic spores and pollen from New Zealand. Palaeont. Bull., Wellington, 22. , 1960: New Zealand Mesozoic and Cainozoic plant microfossils. Palaeont. Bull. Wellington, 32. DEANE, H . , 1904: Further notes on the Cainozoic flora of Sentinel Rock, Otway Coast. Rec. geol. Surv. Vict., 1, pp. 212-215. DORMAN, F. H . , 1966. Australian Tertiary paleotemperatures. J. Geol, 74, pp. 4 9 - 6 1 . , & GILL, E. D., 1959: Oxygen isotope palaeotemperature measurements on Australian fossils. Proc. R. Soc. Vict., 71, pp. 73-98. FISER, JUDITH, & WALKER, D . , 1 9 6 7 : Notes on the pollen morphology of Drimys Forst., section Tasmannia (R. Br.) F. Muell. Pollen Spores,

HARRIS, WAYNE K., 1971: Tertiary

stratigraphic palynology, Otway Basin; in Wopfner, H., & Douglas, J. G. (Eds), The Otway Basin of southeastern Australia. Spec. Bull. geol. Survs S. Aust. & Vict., pp. 67-85. KOHLER, E., 1 9 6 5 : Die Pollenmorphologie der biovulaten Euphorbiaceae und ihre Bedeutung fur die Taxonomie. Grana palynol., 6, pp. 26-120. MARTIN, HELENE A., 1 9 6 9 : The palynology of some Tertiary and later deposits in New South Wales. Ph.D. Thesis, Univ. N.S.W. [unpublished]. , (in press): The palynology of some Tertiary and Pleistocene deposits, Lachlan River Valley, New South Wales. Aust. J. Bot., Supp. Ser. 6. MCWHAE, KATHLEEN M., 1 9 5 7 : A note on the pollen of Whiteodendren and Kjellbergiodendron (Myrtacae). Reinwardia, 4, pp. 189-191. NILSSON, O. 1967: Studies in Montia L . and Claytonia L. and allied genera. III. Pollen morphology. Grana palynol., 7, pp. 279-363. PARTRIDGE, A. D . , 1 9 7 1 : Stratigraphic palynology of the onshore Tertiary sediments of the Gippsland Basin, Victoria. M.Sc. Thesis, Univ. N.S.W. [unpublished]. PATTON, R . T . , 1 9 2 8 : Fossil plants of the Stony Creek Basin. Proc. R. Soc. Vict., 40, pp. 88-90. PELS, S., 1960:

The geology of the Murrumbidgee Irrigation area and surrounding districts. Bull. Water Cons. & Irrig. Comm. N.S.W., 5. WARDLE, P., 1964: Facets of the distribution of forest vegetation in New Zealand. N.Z. Jl Bot., 2, pp. 352-366. 9, p p . 2 2 9 - 2 3 9 . WILLIAMSON, W. H., 1964: The development of ground-water resources of alluvial formations. FRAZER, LILIAN, & VICKERY, JOYCE W., 1 9 3 8 : T h e Bll in Water Resources, Use and Manageecology of the Upper Williams River and ment. Proceedings of Symposium, Australian Barrington Tops districts. II. The rainforest Academy of Science, Melbourne University formations. Proc. Linn. Soc. N.S.W., 63, pp. Press. 139-184. Helene A. Martin, School of Botany, University of New South Wales, Box 1, P.O., Kensington, N.S.W. 2033.

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W. 51 APPENDIX The fossil pollen taxa which have not been of the localities discussed, associated with other previously described are presented in this section. gymnosperms. The holotypes of new species are lodged with ANGIOSPERMAE—DICOTYLEDONAE the Australian Museum. The location of the holotype refers to the microscope BS/4369/M/GB Drimys sp. and a master slide is included with the type slides. (Plate 1, Figs 7, 8) The names of new species not yet published (MarDescription: The pollen occurs in tetrads, intin, in press), cannot be used in this paper so dividual grains (12 (18) 22M- diameter) spherical they are allotted a code number. to sub-oblate with the sides flattened where they touch in the tetrad. The free distal pole has a PTERIDOPHYTA single large round, oval or irregular pore, 6 (8) Cyatheacidites annulata Cookson, 1947 14pt in diameter. The exine, 0.5-IM- thick, is coarsely reticulate over the distal surface, except (Plate 1, Figs 1-3) Comments: Specimens of this species have been for the pore. The muri (2-3pi high, 0.5M, wide) found only at Wagga, in association with Notho- are wavy and beaded, the lumina irregularly polyfagus and gymnosperms. This species is more gonal (3-4M- in diameter) with 2-3 rows of commonly found in the Lower Tertiary but Cook- lumina between the pore and the adjoining grain son (1957) records one probable Pliocene occur- of the tetrad. The reticulum forms a ragged edge around the pore. rence from Queensland. Size: 22 (27) 30m diameter of tetrad. Form B-17 (new species, Martin, in press) Affinities: This fossil Drimys is very similar to (Plate 1, Figs 4, 5) the extant D. insipida (R. Br. ex D.C.) Pilger Description: The spore is piano- to slightly (Fiser & Walker, 1967) and D. lanceolata (Poir.) pyramidal-convex; amb triangular with straight Baill. (PL 1, Fig. 9). The former is slightly larger, sides and broadly rounded angles. The simple tri- corresponding to the larger fossil specimens, lete laesurae extend almost to the equator, the whereas the latter corresponds to the average of margo is very narrow and slightly raised. The the fossils. Pseudowintera axillaris (J. R. & G. exine (3-4^) has a nexine of about IJLX and Forst.) Dandy is much larger (average 39M-), has sexine of 2-3M-, which is thicker around the corners. a larger reticulum with coarser beading and less The pattern is reticulate, with muri 1-1.5M- wide wavy muri when compared with the fossil. and lumina 2-6jx. The pattern covers both proxiDistribution: Lachlan River Valley, Wagga, and mal and distal surfaces but it is smaller and thin- Narrandera. ner on the former. Portulacaceae gen. et sp. indet. Size: 45 (49) 56M- (5 specimens only), equa(Plate 1, Figs 10, 11) torial diameter. Description: The grains are 15-pantocolpate with Comments: The above description applies par- the colpi arranged to form 2 pentagons on oppoticularly to the Lachlan River assemblage in site sides and 5 squares or rectangles between the which the species is rare. At Wagga, where it is The exine is tectate, about 2\i thick, more common, there is a greater variation and pentagons. the nexine thin and the sexine baculate and some specimens have wider muri, especially on finely reticulate. the distal surface. Size: 38M- (one specimen only) largest dimenDistribution: Rare in the Lachlan River Valley, sion. more common at Wagga. Affinities: This fossil can be placed in the Montia pollen type of the Portulacaceae (Nilsson, GYMNOSPERMAE 1967). Four genera within the subfamily MontioCupressaceae gen. et sp. indet. ideae have this pollen type. It occurs also in Calandrinia of the Portulacoideae, but Nilsson (Plate 1, Fig. 6) {op. cit.) has not found this type of pollen Description: The grains are presumed to be amongst Australian species that he has exoriginally spherical, now invariably split, with amined. Athespecimen Montia fontana L. yielded folds typically parallel to the split. There is 12-pantocolpate grainsof with two squares at oppousually some crumpling and an aperture cannot site ends and 4 rectangles (see Figs 12, 13). be detected. The exine is thin, from less than However, there is appreciable variation within the 0.5M- to 0.7M-, and covered with irregularly pollen type and Nilsson {op. cit.) a tetarranged granules which are seen only on well- raploid of Montia fontana which describes is 15-pantocolpreserved specimens. pate whereas the diploid is 12-pantocolpate. Size: 21 (26) 30jx (6 specimens only), greatest Montia fontana is found in southeastern Ausdimension. tralia today. Distribution: Present in low frequencies at each Distribution: Rare, found only at Narrandera. Spec.Publs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

HELENE A. MARTIN Size: 13 (14) 15^t polar diameter x 9 (10) Quintinia sp. \2\i equatorial diameter (5 specimens only). (Plate 1, Figs 14, 15) Affinities: The size and form are like some Description: The 4-5 colpate grain is sub- species of the Cunoniaceae. spherical to prolate; amb circular. It has long, Distribution: Lachlan River Valley, Narransimple colpi which end with a small semicircular dera, and Hay, found only in association with expansion. There may be weak, inconspicuous pores associated with the colpi. The exine (l-1.5n Nothofagus. thick) becomes thinner towards the colpi when Form R-28 (new species in Martin, in press) seen in equatorial outline. It is distinctly twolayered, and is either very faintly patterned or (Plate 2, Figs 6-8) almost psilate. Description: The pollen grains are oblate to Size: 12 (13) 15\i polar diameter x 7 (10) suboblate; amb circular, and there are 5 or 6 short \2\i equatorial diameter (11 specimens), 16,u, colpi, 3-4^ long and lpi wide when expanded. polar view (one specimen only). The exine (1.5-2n thick) is conspicuously twoAffinities: The fossil compares very favourably layered, the layers of equal thickness. The outer with Quintinia sp. (PL 1, Figs 16, 17) which has layer appears structureless and the inner finely 4-6 colpi ending in small, semicircular expansions structured. The outer layer becomes thinner toand weak, poorly defined pores. Like the fossil wards the colpi which appear slightly sunken in grains, these have a faintly patterned exine which polar view. There is a very fine, faint surface is distinctly two-layered, becoming thinner to- pattern, reticulate or more often indeterminate. wards the pores. The size range of both is pracSize: 14 (17) 18jLt, equatorial diameter, 15^ tically identical. polar diameter (the latter one measurement only). Distribution: Found in each locality, only in Distribution: Found at each locality, in assoassociation with Nothofagus. ciation with the Myrtaceae-Casuarina phase. Micrantheum sp. Form Q-32 (new species in Martin, in press) (Plate 2, Figs 1, 2) (Plate 2, Figs 9, 10) Description: The spherical pollen grains have Description: The tricolporate pollen grains are many (45-55) small pores (1-2M« in diam.) occa- oblate with a three-lobed outline when fully exsionally with a narrow border, and polygonal panded, less frequently prolate when contracted markings may be visible between the pores. The and folded. The colpus has a wide smooth memexine is 1-3^ thick, rigid, covered with spines, 1.5- brane, 3|i at the equator where it is interrupted 3|x long and no more than 1|LI wide at the base. The by a characteristic cross colpus, about 8jj, long spines are usually curved, those around the and which forms a solid, transverse border to the pores curving outwards, like a rosette. There may pore. The exine, 1.5-2^ thick, has a reticulum be a faint granular pattern on the exine between with lumina 0.5p, wide, clavate muri In high and 0.5M- wide at the surface. The reticulum is inthe spines. terrupted at the transverse border which appears Size: 19 (24) 29\i largest dimension. solid. An occasional specimen is tectate, with a Affinities: This fossil species is very like Micran- thicker exine. theum ericoides Des. f. (Erdtman, 1952) and M. Equatorial view; 18 (22) 26^ polar axis hexandrum Hook. f. (Plate 2, Fig. 3 and Kohler, x Size: 13 (16) 22^ equatorial axis. Polar view: 18 1965). The two modern species are slightly larger, with an average of about 29\i compared with 24\i, (23) 27^. Distribution: Found at each locality, in associafor the fossil. with the Myrtaccae-Casuarina phase and less Distribution: Lachlan River Valley, Wagga, and tion frequently with gymnosperms. Narrandera, in association with the MyrtaceaeCasuarina phase of the Pliocene. Tricolpites geranioides Couper, 1960 (Plate 2, Figs 11-13) Form P-30 (new species, Martin, in press) Size: 50-60^ polar view (2 specimens only). (Plate 2, Figs 4, 5) Comments: These specimens compare well with Description: The tricolpate grains are prolate with simple, long colpi extending to the poles. Couper's description except that the lumina are The exine (1-1.5m- wide) has a thin nexine and smaller (1-2M- compared with 3-4n for the latter). thick sexine which is baculate and reticulate This distinctive structure of the sexine is found (lumina about 0.5^; muri narrow, < 0.5^). on Geranium spp. (see Plate 2, Figs 14-16) which, Most grains are intectate but occasionally a tec- however, are tricolporate with an extremely tate specimen is seen and then the exine is large pore occupying most of the length of the thicker. The reticulum is usually well defined, colpus. The part of the colpus extending beyond the pore is indistinct and not easily detectable less clearly seen on the tectate grains. 52

UPPER TERTIARY PALYNOLOGY IN SOUTHERN N.S.W. 53 amongst the ornamentation. In polar view, these junctions of the muri, i.e., in the corners of the fossils have an outline similar to Geranium, sug- lumina. The grains are often broken. gesting that they may also be tricolporate, but this Size: 39-44 polar diameter x 29-40|x equatorial cannot be determined directly from the specimens. diameter (5 specimens only). Distribution: Found only at Narrandera where Holotype: AM 6488/7, 73.7 39.6. Plate 2, Figs it is rare. 17, 18. Type Locality: Narrandera, Bore 25394 at 131-137 ft. Tricolporopollenites pelargonioides sp. nov. Affinities: This fossil is very like Pelargonium (Plate 2, Figs 17, 18) spp. (see Plate 2, Figs 19-21) being tricolporate Diagnosis: The grains are subspherical tricol- with large pores. The detail of the sexine on porate with extremely large pores, 10jx in length Pelargonium is exactly the same as described occupying most of the length of the colpus. The for the fossil. The surface pattern of Pelargonium remainder of the colpus, on either side of the enables it to be distinguished from Geranium pore, is inconspicuous amongst the ornamentation. (see above) and Erodium which has the pila in The exine is about A\i thick, nexine and the longitudinal rows, forming 'finger-print' patterns. sexine (about 3|n) has bacula with small pila Whereas the pila are confined to the junctions of which form a reticulate pattern. The lumina are the muri on Tricolporopollenites pelargonioides, l-3(x in diameter, muri about 0.5|LI which are this feature is not evident on Tricolpites geraquite straight between the points of juncture. nioides Couper, 1960. The most conspicuous feature is the pila (about Distribution: Only at Narrandera where it is In in diameter) which are placed exactly at the infrequent.

Spec.Publs geol.Soc.Aust., 4: pp. 35-54, Pis 1-2, 1973.

54

HELENE A. MARTIN EXPLANATION OF PLATES All photographs are from unretouched negatives.

Figs 1-3.

PLATE 1

Cyatheacidites annulata Cookson, 1947, X600. Fig. 1, proximal focus showing the conspicuous verrucate thickenings. Fig. 2, equatorial focus showing the annular flange. Fig. 3, distal focus showing the pitted sculpture. Wagga, Bore 25356 at 216-225 ft, AM6511/7. Fig. 1, 98.6 34.0. Fig. 2, 84.4 39.3. Fig. 3, 84.2 39.4. Figs 4,5. Form B-17 XI,000. Fig. 4, proximal focus showing laesurae and reticulum. Fig. 5, specimen with sunken proximal surface. Lachlan River Valley, Bore 14747 at 259-261 ft. AM6466. Fig. 4, 85.0 31.5. Fig. 5, 89.4 32.4. Fig. 6. Cupressaceae gen. et sp. indet, XI,000, showing typical splitting and folding. Lachlan River Valley, Bore 14505, AM6462 106.6 36.5. Figs 7,8. Drimys sp., XI,000. Fig. 7, surface focus showing the reticulum. Fig. 8, showing the tetrad and distal pore. Lachlan River Valley, Bore 14747 at 222-223 ft, AM6465 83.7 48.3, both figures. Fig. 9. Drimys lanceolata (Poir.) Baill., Modern pollen. Figs 10, 11. Portulacaceae gen. et sp. indet., XI,000. Fig. 10, optical section of part of the exine. Fig. 11 shows one of the pentagons. Narrandera Bore 25394 at 131-137 ft, AM6488/8 84.5 38.3, both figures. Figs 12, 13. Montia fontana L., modern pollen, XI,000. Fig. 12, focus on the end of the grain, showing one square. Fig. 13, side view, with squares at either end. Figs 14, 15. Quintinia sp. XI,000. Polar and equatorial views respectively. Fig. 14, Lachlan River Valley Bore 14747 at 259-261 ft, AM6466 78.0 32.2. Fig. 15, Ivanhoe Bore 21296 at 290-295 ft, AM6470 91.4 32.9. Figs. 16, 17. Quintinia sp., modern pollen, XI,000. Polar and equatorial views respectively. Figs 1,2.

PLATE 2

Micrantheum sp., XI,000. Fig. 1, surface focus showing pores and surrounding spines. Fig. 2 shows the thick wall in optical section. Lachlan River Valley Bore 14576 at 133-142 ft. AM6484 88.2 48.2, both figures. Fig. 3. Micrantheum hexandrum Hook, f., modern pollen, XI,000. Surface view showing spines. Figs 4 5. Form P-30, XI,000. Fig. 4, surface view showing reticulum. Fig. 5, optical section of exine. Ivanhoe Bore 21296 at 290-295 ft, AM6470 72.7 39.7, both figures. Figs 6-8. Form R-28, XI,000. Fig. 6, optical section; Fig. 7, surface focus; and Fig. 8, equatorial view showing one of the colpi. Lachlan River Valley Bore 14747, Fig. 6 at 159-160 ft. AM6463 85.7 45.5. Fig. 7 at 222-223 ft. AM6465 90.1 32.0. Fig. 8 at 216-218 ft. AM6464 75.2 32.2. Figs 9 10. Form Q-32, XI,000. Fig. 9, oblique polar view showing one colpus and the conspicuously thickened borders of the cross colpus. Fig. 10, equatorial view. Lachlan River Valley Bore 14747 at 159-160 ft, AM6463. Fig. 9, 74.5 44.1 Fig. 10, 102.5 38.2. Figs 11-13. Tricolpites geranioides Couper, 1960. Fig. 11, polar view, X600. Fig. 12, surface focus and Fig. 13, optical section of exine, both XI,000. Narrandera Bore 25394 at 106110 ft. AM6489/6 98.6 48.2, all three figures. Figs 14-16. Geranium spp., modern pollen. Fig. 14, Geranium pilosum Forst. f. ex Willd., polar view, X600. Fig. 15, the same species and Fig. 16, Geranium solandri Carolyn, surface focus; both figures, XI,000. Figs 17, 18. Tricolporopollenites pelargonioides sp. nov., XI,000. Fig. 17, optical section. Arrows indicate the edges of a pore. Fig. 18, surface focus showing the characteristic pattern. Narrandera Bore 25394 at 131-137 ft, AM6488/7 73.7 39.6, both figures. Figs 19-21. Pelargonium spp., modern pollen. Fig. 19, Pelargonium australe Willd., polar view^ and Fig. 20, Pelargonium rodneyanum Lindl., equatorial view showing the extremely large pore, both figures X600. Fig. 21, the latter species, surface focus, XI,000.

HELENE A . MARTIN

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE 1

PLATE 2

HELENE A .

MARTIN

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION, OFFSHORE GIPPSLAND BASIN, AUSTRALIA By LEWIS E. STOVER & P. R I C H A R D EVANS (With 2 Tables, 2 Text-Figures and 4 Plates) ABSTRACT

Recent exploration in the Gippsland Basin afforded the opportunity to examine in considerable detail the spore-pollen assemblages from more than 800 conventional and sidewall cores taken from more than 40 offshore wells. Samples used in this study are from the dominantly non-marine Latrobe Group of Late Cretaceous to Late Eocene age. Within the Latrobe Group, seven palynologic zones are recognized. These, in ascending order are: the Late Cretaceous Nothofagidites senectus and Tricolporites lilliei Zones, the Palaeocene Tricolpites longus and Lygistepollenites balmei Zones and the Eocene Malvacipollis diversus, Proteacidites asperopolus and Nothofagidites asperus Zones. Each of the zones is identified in several wells and the continuity of the biostratigraphic units can be traced across the entire offshore part of the Gippsland Basin. In addition, the vertical succession of zones is seen repeatedly in numerous wells, and the spore-pollen zonation within the Latrobe Group is interpreted as representing an essentially uninterrupted sequence. Associated microplankton from the Palaeocene and Eocene zones provide age-confirming information. INTRODUCTION The Gippsland Basin lies partly onshore but mainly offshore between the coast of eastern Victoria and the Bassian Rise, a basement high defined by geophysical means, which extends between the granitic masses of Wilsons Promontory in Victoria and Flinders Island near Tasmania. The basin encompasses about 48,000 sq. km measured to the 150 m water depth and contains Lower Cretaceous to Recent strata estimated to have a maximum thickness of approximately 7,500 m in the centre of the basin. Structurally, the basin is divided into three major parts (Fig. 1) which from north to south are the North Platform, the central Deep Basin and the South Platform. Onshore exploration for commercial accumulations of hydrocarbons has been active for a number of years and the northwestern onshore part of the basin in the Latrobe Valley contains extensive brown coal deposits utilized for the generation of electricity. Since 1965 the offshore part of the Gippsland Basin has been explored successfully by Esso Australia Ltd and Broken Hill Proprietary Co. Ltd (BHP) with the discovery of significant petroleum and gas reserves (Weeks & Hopkins, 1966; Wallis, 1967; Richards & Hopkins, 1969). During the early stages of offshore exploration it was realized that the major hydrocarbon

accumulations occurred largely in the nonmarine pre-Oligocene rocks, and that spores and pollen would most likely provide a means for dating and correlating Late Cretaceous, Palaeocene and Eocene subsurface sections. Consequently, a study of spore-pollen assemblages from a limited number of wells was undertaken by Dr L. E. Stover at the Esso Production Research Company in Houston, Texas, U.S.A., on behalf of Esso Australia, and at the same time Dr Mary E. Dettmann of the University of Queensland provided palynologic determinations and interpretations for currently drilled wells. In 1968, Esso established a palynologic laboratory in Sydney under the direction of Dr P. R. Evans. Eventually, the spore-pollen zonation proposed earlier by Stover was further documented, refined, and utilized for identifying pre-Oligocene sections in all of BHP-Esso's offshore Bass Strait wells. This paper summarizes the results of these studies where applicable to the Gippsland Basin and concerns only the main characteristics of the spore-pollen assemblages from the Late Cretaceous to Eocene intervals. A more comprehensive taxonomic treatment of the assemblages is planned for publication elsewhere. The zonation presented here is based on the occurrences and distribution of spores and pollen from about 800 conventional and

Spec.Publs geol.Soc.Aust., 4: pp. 55-72, Pis 1-4, 1973.

LEWIS E. STOVER & P. RICHARD EVANS

56

latter group. Oligocene and Miocene strata overlying the Latrobe Group are marine, as evidenced by their excellent planktonic foraminiferal assemblages, and are divided into the Lakes Entrance Formation below and the Gippsland Limestone above. The foraminiferal zonules shown in Figure 2 are those of D. J. Taylor and modified somewhat from his earlier scheme (Taylor, 1966). The spore-pollen zones STRATIGRAPHY the Nothofagidites senectus to the N. The stratigraphic sequence and the lithologic from asperus are the main subject of the next units described by James & Evans (1971) for section. the offshore part of the Gippsland Basin are depicted in Figure 2. The Lower Cretaceous S P O R E - P O L L E N Z O N A T I O N Strzelecki Group is represented by non-marine CRETACEOUS ZONES rocks as is the Upper Cretaceous to Eocene Spore-pollen zones determined by Dettmann Latrobe Group; a few marine strata occur sporadically throughout the upper part of the & Playford (1969) in the Great Artesian and

sidewall core samples from more than 40 wells (Fig. 2). The pre-Oligocene strata penetrated by each of these wells average around 1,200 m. Extensive sample coverage enabled the recognition of the same sequence of assemblages in numerous wells and gave rise to a high level of confidence in the zonation.

ESS0-B.H.R LOCALITY M A P 0

GIPPSLAND BASIN

A B A N D O N E D OIL W E L L

•

*

ABANDONED GAS WELL

^ A B A N 0 0 N E 0 WITH GAS

•

DRILLING P L A T F O R M

^ DRY & A B A N D O N E D W E L L

A B A N D O N E D WITH OIL S H O W SHOW

XNGESp

Halibut-1

19 G r o p e r - 1

25 B r e a m - 2

31 S n a p p e r - 2

8 Kingfish-1

20 M u l l e t - 1

26 F l o u n d e r - 3

32 P l a t f o r m s

9 Dolphin-1

21 S a l m o n - !

27 M a c k e r e l - 1

33 B l u e b o n e - 1

10 K i n g f i s h - 2

22 F l o u n d e r - 2

28 F l a t h e a d - 1

34Groper-2

11 K i n g f i s h - 3

23 P l a t f o r m

29 T u r r u m - 1

35 B a r r a c o u t a - 3

12 P l a t f o r m

24 B r e a m - I

30 W a h o o - 1

38 G u r n a r d - 1

7

Fig. 1. Location map showing basins and wells (10 miles = 16 km). (Modified from James & Evans, 1971).

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION

BIOSTRATIGRAPHIC ZONES

MIOCENE

FORAMS

EPOCH

OLIGOCENE EOCENE PALEOCENE

-SPORE-PI

uu UPPER CRETACEOUS c •3

LOWER CRETACEOUS

N.W.

A B-E

STRATIGRAPHY OFFSHORE GIPPSLAND S.E. PLIOCENE-RECENT Up to 1000'Marine Calcarenite GIPPSLAND LIMESTONE Up to 5000' Marine LAKES ENTRANCE FM. Up to 1500' Marine

F-H H2 h I2-J2 !i.dd[Jtf 1 lib P.asperopolus M.rfiversus L.balmei T.longus T. lilliei

TH

GURNARD FM.

\

|v

TURRUM FM 1

lllllllllllllllllllll >FLOUNDER FM.

w

LATROBE GP \ \ \

N. senectus T.pachyexinus C.triplex A.distocarioatus T. pannosus C.paradoxa

57

i l k

Predominantly Non-marine Lacustrine and Fluviatile Estimated up to 15,000'

\ STRZELECKI GROUP Up to 10.000' Non-marine

D.speciosus C.stylosus

Fig. 2. Stratigraphic section. Otway Basins have been recognized in only a few Gippsland Basin wells. Because strata older than those containing the Late Cretaceous Nothofagidites Microflora of Dettmann & Playford are rarely encountered by drilling in the Basin Deep, the following discussion is confined to spore-pollen assemblages and zones considered equivalent for the most part to the Nothofagidites Microflora. The youngest Late Cretaceous assemblage identified in the Otway Basin by Dettmann & Playford is the Nothofagidites Microflora. By labelling it a microflora they were apparently unprepared at that time to designate formally the interval with the assemblage as a zone. They recognized correctly that they were dealing with an assemblage that is older than the Palaeocene Triorites edwardsii Assemblage of Harris (1965) from the Pebble Point Formation in south-

western Victorian coastal sections. Because a regional unconformity occurs between the Sherbrook Group with the Nothofagidites Microflora and the Pebble Point Formation with the Triorites edwardsii Assemblage, no means are available in the Otway Basin for ascertaining how the Nothofagidites Microflora is replaced by the Triorites edwardsii Assemblage. Spore-pollen assemblages from the Gippsland Basin as well as additional information from some offshore Otway Basin wells indicate that the interval through which the Nothofagidites Microflora ranges is divisible into two zones, here named the Nothofagidites senectus Zone and the Tricolporites lilliei Zone. In the Gippsland Basin, additional zones of Palaeocene age overlie the T. lilliei Zone, thus a fossiliferous interval, not present in the

Spec.Publs geol.Soc.Aust, 4: pp. 55-72, Pis 1-4, 1973.

58

LEWIS

E. S T O V E R

& P.

Otway Basin, and lying between the Nothofagidites Microflora and Triorites edwardsii Assemblage equivalents, occurs in the Gippsland Basin. Within this basin, an uninterrupted sequence of spore-pollen assemblages is identifiable from the Upper Cretaceous through the Palaeocene. Nothofagidites senectus Zone. The base of the zone is defined by the first appearance of specimens of Nothofagidites spp., including the nominate species, N. senectus Dettmann & Playford, 1968; fossil specimens of Nothofagidites spp. are referable to the Nothofagus brassi Group of extant forms. Specimens assignable to the N. fusca or N. menziesii Groups have as yet not been identified from this part of the section. The top of the zone is marked by the first appearance of Tricolporites lilliei (Couper) Stover & Evans, 1973. Triorites edwardsii Cookson & Pike, 1954, Tricolpites gillii Cookson, 1957, T. sabulosus Dettmann & Playford, 1968 and Proteacidites amolosexinus Dettmann & Playford, 1968 have their first appearance in the N. senectus Zone. Most of the triporate pollen from this zone, apart from P. amolosexinus, have relatively simple wall structures. Periporate and stephanoporate pollen, such as Caryophyllidites polyoratus Couper, 1960 and Australopollis obscurus (Harris) Krutzsch, 1966, occur consistently and trilete spores, monolete spores, and gymnosperm pollen are abundant. The Nothofagidites senectus Zone is equivalent at least in part to the lower half of the Nothofagidites Microflora. With the exception of Stereisporites viriosus Dettmann & Playford 1968, all of species reported from the lower part of the Nothofagidites Microflora in the Otway Basin have been identified in the N. senectus Zone in the Gippsland Basin. The zone has been identified mainly in the marginal wells of the basin, hence no meaningful estimates of its average thickness can be made and no section in which the zone is characteristically developed can be designated at this time. Tricolporites lilliei Zone. This zone is characterized by the first appearance of Tricolporites lilliei at the base of the zone, by a marked increase in the relative abundance of Nothofagidites spp., and by the introduction of several pollen and spore species. Many of latter represent undescribed forms such as Triporopollenites sp. A. Described species which make their first appearance in the T. lilliei Zone include Tricolpites waiparaensis

RICHARD

EVANS

Couper, 1960, Stereisporites regium (Drozh.) Drugg, 1967, Phyllocladidites verrucosus (Cookson) Stover & Evans, 1973, Lygistepollenites florinii (Cookson & Pike) Stover & Evans, 1973, Proteacidites palisadus Couper, 1953, Nothofagidites endurus Stover & Evans, 1973. Tricolpites sabulosus and Nothofagidites senectus terminate in the T. lilliei Zone. Not only is there an increase in the number of specimens and species of Nothofagidites, all of the Nothofagus brassi Group, in the J. lilliei Zone, but also a greater diversity among other types of angiosperm pollen. There is also a decided tendency towards the introduction of pollen with more complex exinal sculpture than found among pollen from the Nothofagidites senectus Zone. The T. lilliei Zone is developed typically in the Barracouta-1 well at 2,653.5 m and occurs also in the Barracouta A-3 platform well between 2,300.9 and 2,449.3 m; it has been observed in several other offshore Gippsland Basin wells. Observed thicknesses for the zone are from 90-180 m. PALAEOCENE

ZONES

Tricolpites longus Zone. The base of the zone is marked by the virtual absence of Nothofagidites spp., by a significant increase in the abundance of Triorites edwardsii, and by the first appearance of Latrobosporites crassus Harris, 1965, Stereisporites (Tripunctisporis) spp., Dilwynites granulatus Harris, 1965, and Proteacidites sp. A. Although not restricted to the T. longus Zone, the unusual planar tetrad (Plate 2, Fig. 4) and Tetracolporites sp. A occur fairly frequently in assemblages from this zone. Several species terminate at or near the top of the zone and these include the nominate species, Tricolpites longus Stover & Evans, 1973, together with Proteacidites palisadus, Tricolporites lilliei, Tricolpites sp. A. Proteacidites amolosexinus does not appear to extend through the full range of the zone, whereas Ornamentifera sentosa Dettmann & Playford, 1968 is found sporadically throughout the zone and extends into the base of the overlying zone. Angiosperm pollen, gymnosperm pollen and spores are common and no one group consistently dominates the assemblages from the T. longus Zone, although gymnosperm pollen may do so locally. The zone averages between 280 and 320 m in thickness and is excellently developed in the Barracouta-1 between 2,345.2 and 2,642.0 m. Lygistepollenites balmei Zone. This zone is

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION defined by the combined occurrence of: 1, a group of species which appear in older zones but do not extend into the overlying zones; 2, another group of species which make their first appearance in the Lygistepollenites balmei Zone; and 3, a third and smaller group of species which occur only within the zone. In the first group are included Lygistepollenites balmei (Cookson) Stover & Evans, 1973, Phyllocladidites verrucosus, Australopollis obscurus and Triorites edwardsii. Triorites harrisii Couper, 1953, Nothofagidites flemingii (Couper) Potonie, 1960, Proteacidites annularis Cookson, 1950, Myrtaceidites parvus Cookson & Pike, 1954 and Trilites tuberculiformis Cookson ex Couper, 1953 are representative of the second group. Species which occur just within this zone are Lygistepollenites ellipticus (Harris) Stover & Evans, 1973, Phyllocladidites reticulosaccatus Harris, 1965, Polycolpites sp. A and possibly Peromonolites densus Harris, 1965. In general, the assemblages from the Lygistepollenites balmei Zone contain a greater number of angiosperm pollen species than those from the underlying Tricolpites longus Zone. The relative abundances of gymnosperm pollen and trilete spores tend to increase in this zone, even though there is no marked increase in the number of species representing these palynomorph groups. Proteaceous pollen are neither particularly diverse nor particularly common, nor are they morphologically complex. Perhaps the most important events from a botanical viewpoint are the introduction of representatives of the Nothofagus fusca Group of southern beech pollen and of myrtaceous pollen in the L. balmei Zone. In so far as can be determined the top of the zone corresponds to the Palaeocene— Eocene boundary. The Lygistepollenites balmei Zone extends, on the average, through 400 to 500 m of the Latrobe Group and the zone has been identified in a majority of the offshore Gippsland Basin wells. Representative sporepollen assemblages from the L. balmei Zone occur, for example, in the Barracouta-1 well between 1,727.2 and 2,211.5 m, in the Marlin2 well between 2,256.6 and 2,653.5 m, and in the Cod-1 well between 2,243.5 and 3,001.8 m. EOCENE ZONES

Malvacipollis diversus Zone. Unlike the preceding zones, angiosperm pollen dominate this assemblage and within the limits of the Malvacipollis diversus Zone a comparatively

59

large number of triporate and tricolporate pollen appear. The lower part of this zone contains a rather meagre assemblage composed mainly of carry-overs from older zones plus a few species which occur for the first time in the Latrobe Group. The latter include Cupanieidites orthoteichus Cookson & Pike, 1954, Spinizonocolpites prominatus (Mclntyre) Stover & Evans, 1973, Tiliaepollenites notabilis Harris, 1965 and Proteacidites dilwynensis Harris, 1965. The nominate species, Malvacipollis diversus Harris, 1965, is usually common in the lower part of the zone, and locally Myrtaceidites parvus may be abundant. There is a gradual increase in the number of angiosperm pollen in the M. diversus Zone owing to the continual introduction of new forms upsection. This, coupled with the persistence of older-occurring species contributes to the distinctness and rich variety of the spore-pollen assemblages from this zone. Some species which appear first in the Malvacipollis diversus Zone are: Santalumidites cainozoicus Cookson & Pike, 1954, Beaupreaidites elegansiformis Cookson, 1950, Proteacidites grandis Cookson, 1950, P. latrobensis Harris, 1966, P. ornatus Harris, 1965, and P. pachypolus Cookson & Pike, 1954. Myrtaceidites tenuis Harris, 1965, occurs only in this zone in the Gippsland Basin. One of the most characteristic features of assemblages from this zone is the prevalence and the large number of species of Proteacidites which are diverse in size, shape and exine sculpture, although many species have some type of a reticulate exine. By the time the top of the zone is reached, the relative abundance of angiosperm pollen commonly exceeds 75 per cent. Proteaceous pollen are certainly the more conspicuous forms and specimens of Nothofagidites spp. of both the Nothofagus brassi and N. fusca Groups, although not rare, are definitely subordinate. Spores are usually rare or sparse as are gymnosperm pollen. The Malvacipollis diversus Zone averages 250-300 m in thickness and is, therefore, generally thinner than the underlying Lygistepollenites balmei Zone. Typical M. diversus spore-pollen assemblages have been recovered from the Marlin-1 well between 1,451.8 and 1,575.9 m and an unusually thick section was identified in the Cod-1 well between 1,947.0 and 2,387.9 m. This zone is widespread in the Gippsland Basin and has been recognized in practically every offshore well.

Spec.Publs geol.Soc.Aust., 4: pp. 55-72, Pis 1-4, 1973.

60

LEWIS E. STOVER & P. RICHARD EVANS

Proteacidites asperopolus Zone. In comparison to the Malvacipolis diversus Zone, the Proteacidites asperopolus Zone is thin and rarely exceeds 75-100 m in thickness. Sporepollen assemblages within this interval are characterised by the first occurrence of Nothofagidites asper us (Cookson) Stover & Evans, 1973 and N. goniatus (Cookson) Stover & Evans, 1973. The presence of these species represents the introduction of the third type of Nothofagus pollen, the N. menziesii Group, into the stratigraphic sequence in the Gippsland Basin. The zone is characterized further by the high abundance of Proteacidites asperopolus Stover & Evans, 1973, or of P. pachypolus, or in some samples, of both species. Also indicative of the P. asperopolus Zone is the greater abundance of specimens of Triorites harrisii over those of Nothofagidites spp. The significance of this becomes evident later. A few species with generally short ranges appear in this zone, for example, Liliacidites sp. A, Monosulcites sp. A and Tricolpites sp. B. These species are illustrated in Plate 4. With the exception of Proteacidites asperopolus and P. pachypolus, which are especially prevalent, several other species of Proteacidites occur less frequently or terminate in the P. asperopolus Zone, as do a few species of other angiosperm genera. Examples of species that do not range above the P. asperopolus Zone in the Gippsland Basin include Tiliaepollenites notabilis, Proteacidites dilwynensis, P. ornatus Harris, 1965 and Spinizonocolpites prominatus. Nothofagidites deminutus (Cookson) Stover & Evans, 1973 is usually common in this zone. Typical spore-pollen assemblages from the P. asperopolus Zone occur in the Marlin-3 well in the conventional core from 1,545.5 to 1,551.8 m. Nothofagidites asperus Zone. The base of the Nothofagidites asperus Zone is indicated by the sudden and dramatic influx of specimens of Nothofagidites spp. into the spore-pollen assemblages. Because of this, there is a sharp reversal in the relative abundances of Nothofagidites spp. and Triorites harrisii. As stated above, in the Proteacidites asperopolus Zone, T. harrisii was more abundant than Nothofagidites spp.; in the N. asperus Zone the reverse is true. Within the lower part of the Nothofagidites asperus Zone the spore-pollen assemblages are still quite diverse, but gradually species drop out and there is only partial replacement through the introduction of new taxa. There

is, then, in the offshore Gippsland Basin a progressive diminution in the diversity of species in the palynomorph assemblages with decreasing geologic time. There is also a concomitant increase in the amount of microplankton. The decrease in the spores and pollen and the increase in the microplankton reflects the spreading of marginal marine conditions across the Gippsland Basin towards the close of Eocene time. Pollen that make their first appearance in the Nothofagidites asperus Zone include Triorites magnificus Cookson, 1950, Tricolpites thomasii Cookson & Pike, 1954, Nothofagidites falcatus (Cookson) Stover & Evans, 1973 as well as Aglaoreidia sp. A and a new species of Myrtaceidites. The Nothofagidites asperus Zone, which occurs over a large part of the Gippsland Basin and is found in the upper 75 to about 250 m of the Latrobe Group, contains few species of Proteacidites. Those that do occur are represented more often than not by a low number of specimens. The zone is well expressed in the Groper-1 well between 931.8 and 1,011.2 m and in the Turrum-1 well between 1,954.7 and 2,037.4 m. The top of the N. asperus Zone cannot be defined palynologically in the Gippsland Basin where it is overlain by post-Eocene marine strata with well-developed planktonic foraminiferal assemblages. AGES OF SPORE-POLLEN ZONES Independent, definitive, age-significant palaeontologic data are sparse in the Latrobe Group of the Gippsland Basin, especially in the interval from the Nothofagidites senectus through the Lygistepollenites balmei Zones, that is, in the Late Cretaceous and Palaeocene. Consequently, comparisons are made with other areas having spore-pollen assemblages in association with other fossil groups such as Foraminifera and microplankton. In the Otway Basin Nautilus-1 well, Taylor {in Lunt & James, 1968) reported late Turonian-early Senonian planktonic Foraminifera in association with spores and pollen suggestive of the N. senectus Zone. It is possible that the base of the N. senectus Zone may coincide with or be very close to the Turonian-Senonian boundary. The base of the zone is marked by the introduction of Nothofagidites spp. and in New Zealand the earliest record of Nothofagidites pollen is from the early Senonian (Arowhanan) from which Couper (1960)

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION 61 identified N. kaitangata Te Punga, 1948. It may pollenites balmei Zone and which compare also be that Nothofagidites pollen appear some- favourably with species from the Danian of what later in the geological record in Aus- California, U.S.A. (Drugg, 1967; Stover, tralia than in New Zealand, and that the base 1973). Thus we regard the Tricolpites longus of the N. senectus Zone could be younger, Zone as early as possibly Middle Palaeocene. and perhaps closer to the Coniacian-Santonian Middle Palaeocene planktonic Foraminifera boundary than to the Turonian-Coniacian were reported from the shell bed within the boundary. There is little doubt, however, that Pebble Point Formation approximately 12 m the N. senectus Zone is Late Cretaceous above the base of the formation (McGowran, (Senonian). 1965), and Cookson (19656) and Cookson & Eisenack (1965c) recorded microplankton The Nothofagidites senectus Zone is suc- from the same strata as well as from samples ceeded with no apparent interruption by the taken at 1.0 to 1.3 m above the base of the Tricolporites lilliei Zone. Evidence for the formation. These Otway Basin samples and Senonian-Maestrichtian age of the latter zone others from the Pebble Point Formation conis that given by Dettmann & Playford (1969) tain typical Lygistepollenites balmei sporefor their Nothofagidites Microflora. In the pollen assemblages (Harris, 1965). Among Otway Basin, dinophyceates in the T. lilliei the microplankton from the Pebble Point ForZone include a form similar to Deflandrea mation, Svalbardella australina Cookson, 1965, korojongensis Cookson & Eisenack (1958), Cyclonephelium retiintextum Cookson, 1965, who reported this species from the Campanian and Deflandrea dilwynensis Cookson&Eisenack, to Maestrichtian of Western Australia, to- 1965 occur in the L. balmei Zone in the Gippsgether with D. pellucida (Deflandre & Cook- land Basin. The first occurrence of Wetzeliella son) Cookson & Eisenack. This combination homomorpha Deflandre & Cookson, 1955 is in of species lends credence to the Late Cre- the upper part of the L. balmei Zone and in taceous age for the T. lilliei Zone. Admittedly, New Zealand its first occurrence is in the convincing data for interpreting more pre- Teurian (Wilson, 1967), which he shows as cisely the ages of the Nothofagidites senectus being Palaeocene. Collectively, foraminiferal and T. lilliei Zones are lacking, and until and microplankton evidence, although limited, more definitive information becomes available leads to the conclusion that the L. balmei Zone it is perhaps best to consider the zones as is Middle to Late Palaeocene. they are developed in the Gippsland Basin as The Lygistepollenites balmei Zonz-Malvasimply Late Cretaceous. Evidence for determining the position of the cipollis diversus Zone boundary coincides with Mesozoic-Tertiary boundary is inconclusive in the Palaeocene-Eocene boundary. This also the absence of firm micropalaeontologic data. corresponds to the Pseudohastigerina datum of Our best approximation of this horizon is at Berggren (1969), which in the Otway Basin the Tricolporites lilliei Zone—Tricolpites correlates with the Rivernook 'A' planktonic longus Zone boundary, based on the decided foraminiferal assemblage of the Dilwyn Clay decrease to virtual absence of specimens of (Taylor, pers. comm.). Microplankton occur Nothofagidites spp. at the base of the T. sporadically throughout the section in the longus Zone. The same situation occurs in Gippsland Basin extending from the base of New Zealand with the marked reduction in the M. diversus to the top of the Proteacidites Nothofagidites spp. in the lower Teurian asperopolus Zone. These dinophyceate assem(Palaeocene) (Couper, 1960). More recently, blages contain many of the species described Jenkins (1971) presented foraminiferal evi- from the Rivernook Member of the Dilwyn dence which indicates that the Teurian section Clay, from the Dartmoor Formation near from which Couper obtained spores and pollen Casterton, Victoria, and from Strahan, Tas(Couper's sample S68/770) is equivalent to mania (Cookson & Eisenack, 19676, 19656, the upper part of the Danian. The other sam- 1967a, respectively). Samples from these ple (S68/768), which Couper shows as being localities also contain associated spores and in the Teurian, is approximately 61 m below pollen indicative of the M. diversus Zone as sample S68/770 and apparently lacks Fora- developed in the Gippsland Basin. Some of the minifera. Spore-pollen assemblages from the more important dinophyceate species described Gippsland Basin contain some microplankton by Cookson & Eisenack include Deflandrea among which are species of Deflandrea that dartmooria, Homotryblium tasmaniense, Kendo not extend into the overlying Lygiste- leyia fimbriata, Wetzeliella hyperacantha and Spec.Publs geoI.Soc.Aust, 4: pp. 55-72, Pis 1-4, 1973.

62

LEWIS E. STOVER & P. RICHARD EVANS

Cordosphaeridium bipolare, all of which occur in the Gippsland Basin. Specimens of Wetzeliella coleothrypta Williams & Downie (in Davey et al., 1966) and W. edwardsii Wilson, 1967 occur in assemblages from the Proteacidites asperopolus Zone. Wilson (1967) reported these species from the Mangaorapan and Heretaungan (middle and late Early Eocene Globorotalia crater crater Zone) in New Zealand. He also shows the two species first occurring together and then with W. edwardsii extending beyond the range of W. coleothrypta, with neither species ranging into the Middle Eocene Porangan. The identical situation has been observed among specimens from the Gippsland Basin. Primarily on this basis, we place the Early Middle Eocene boundary at the top of the P. asperopolus Zone and interpret the Malvacipollis diversus and P. asperopolus Zones as early Eocene. The Middle to Late Eocene age for the Nothofagidites asperus Zone is based on associated microplankton and the occasional occurrence of planktonic Foraminifera, for example, Globigerina linaperta Finlay and G. ampliapertura Bolli. Dinophyceate assemblages from the N. asperus Zone are quite similar to those described from the Otway Basin Browns Creek Clays section near the Johanna River in southwestern Victoria (Cookson, 1965a; Cookson & Eisenack, 1965a). The following species, all known from the Browns Creek Clays, have been observed in the N. asperus Zone in the Gippsland Basin: Deflandrea phosphoritica Eisenack, 1938, Deflandrea heterophlycta Deflandre & Cookson, 1955, Gymnodinium australiense Deflandre & Cookson, 1955, Leptodinium dispertitum Cookson & Eisenack, 1965, Eisenackia ornata Cookson & Eisenack, 1965, Schematophora speciosa Deflandre & Cookson, 1955, Systematophora ancyrea Cookson & Eisenack, 1965, Cordosphaeridium capricornum Cookson & Eisenack, 1965 and Hystrichokolpoma rigaudae Deflandre & Cookson, 1955. From the same section Taylor obtained rich planktonic foraminiferal assemblages upon which he based his Middle Eocene N Zonule and Late Eocene M Zonule (Taylor, pers. comm.). Taken in reverse order, collaborative palaeontologic data afforded by microplankton and Foraminifera provide a firm Eocene age assignment for the sections in the Gippsland Basin containing the Nothofagidites asperus, Proteacidites asperopolus and Malvacipollis

diversus spore-pollen zones. The Palaeocene age for the Lygistepollenites balmei and Tricolpites longus Zones is less well documented, especially the latter whereas concrete evidence for interpreting the ages of the Tricolporites lilliei and Nothofagidites senectus Zones more precisely than Late Cretaceous is not yet available. SUMMARY Petroleum exploration in the offshore Gippsland Basin and the discovery of substantial hydrocarbon reserves provided material and impetus for studying palynomorphs most useful for dating, sub-dividing and correlating Late Cretaceous through Eocene subsurface sections. Within the approximately 1,3001,700 m of dominantly clastic, non-marine Latrobe Group strata commonly penetrated in offshore wells, 7 spore-pollen zones are recognized. The Late Cretaceous part of the section contains the Nothofagidites senectus and the Tricolporites lilliei Zones; the Palaeocene part has the Tricolpites longus and Lygistepollenites balmei Zones and the Eocene interval includes the Malvacipollis diversus, Proteacidites asperopolus and Nothofagidites asperus Zones. These zones represent a continuous, gradually changing progression of spore-pollen assemblages as depicted in part by the stratigraphic chart (Table I), which shows the ranges of the spore and pollen species cited in the text and/or illustrated on Plates 1-4. In defining and selecting zone limits, such factors were taken into account as 1, the first and last occurrence of species, 2, the total composition of the assemblage and 3, the relative abundances of certain forms. Emphasis was placed on the overall character of the assemblage rather than on the ranges of a few selected species, although ranges were certainly considered in defining zone boundaries. This approach enabled zones to be identified even though some of the species known to occur in the zone may be lacking in some samples. This method is practical, reliable and effective for regional correlation across the entire offshore part of the Gippsland Basin. Because of the large number of core samples examined and the density of wells in the Gippsland Basin, the lateral continuity of zones can be clearly demonstrated and the same vertical succession of spore-pollen zones is seen repeatedly in well after well. There is no evidence of a major palynologic break from the Late Cretaceous to the top of the Eocene section,

>-ATE

EARLY TO MIDDLE |

CRETACEOUS MIDDLE TO LATE

PALAEOCENE EARLY

EOCENE I

MIDDLE TO LATE

AGE ZONES

ohaiensis

gillii polyoratus

lilliei

regium

NOTHOFAGIDITES

ASPERUS

PROTEACIDITES

ASPEROPOLUS

MALVACIPOLLIS

DIVERSUS

LYGISTEPOLLENITES

BALMEI

TRICOLPITES

LONGUS

TRICOLPORITES LILLIEI

NOTHOFAGIDITES

SENECTUS

Table 1. Species distribution chart

sp. A

sp. A

sp. A

. Malvacipollis

sp. A tenuis

> Ischyosporites Myrtaceidites

cainozoicus pachypolus

waterbolkii

deminutus

sp. A

aspersus

sp. B

falcatus

sp. A Nothofagidites

thomasii Aglaoreidia

magnificus Tricolpites

Triorites

Nothofagidites goniatus

Nothofagidites

sp. A

Liliacidites Tricolpites

Tetrahedral tetrad

Tricolporites

Monosulcites s p . A

Nothofagidites

Proteacidites asperopolus

Proteacidites latrobensis

Beaupreaidites elegansiformis

Kuylisporites

Proteacidites grandis

Proteacidites sp. B

Proteacidites

Santalumidites

Proteacidites ornatus

orthoteichus

. Cupanieidites

prominatus

dilwynensis Spinizonocolpites

Proteacidites

notabilis

parvus diversus

. Myrtaceidites

Tiliaepollenites

annularis

- Proteacidites

. Nothofagidites flemingii

Rugulatisporites

. Triorites harrisii

Trilites tuberculiformis

Polycolpites

ellipticus reticulosaccatus

Lygistepollenites Phyllocladidites

Peromonolites densus

spp.

granulatus - S. (Tripunctisporis)

- Dilwynites

crassus

sp. A Latrobosporites

Proteacidites

Nothofagidites endurus

Tetracolporites

Planar tetrad

Proteacidites palisadus

- Lygistepollenites florinii

Phyllocladidites verrucosus

Stereisporites

Tricolpites waiparaensis

Tricolporites

Tricolpites longus

sp. A

balmei

sentosa

Camarozonosporites

Lygistepollenites

Ornamentifera

Triporopollenites sp. A

Tricolpites sp A

Caryophyllidites

Tricolpites

Australopollis obscurus

Triorites edwardsii

Proteacidites amolosexinus

Camarozonosporites

senectus

SPORE -

Nothofagidites

Tricolpites sabulosus

POLLEN

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION or to express it in zone terms, from the Nothofagidites senectus to the N. asperus Zones. The zonal concepts and definitions established for the Gippsland Basin may prove to be equally applicable in the adjacent Bass Basin as well as the Otway Basin. DESCRIPTIVE PALAEONTOLOGY Genus Phyllocladidites Cookson ex Couper, 1953, emend. Type species: Phyllocladidites mawsonii Cookson, 1947, designated by Couper (1953, p. 38.) 1947 Disaccites (Phyllocladidites), Cookson, p. 132. nom. nud. 1953 Dacrydiumites Cookson, p. 66. nom. nud. 1953 Phyllocladidites Cookson ex Couper, p. 36. 1957 Dacrydiumites Cookson, Cookson, p. 46. pars. 1958 Phyllocladidites (Cookson) Couper, Potonie, p. 69. 1960 Dacrydiumites Cookson, Couper, p. 43. 1965 Phyllocladidites Cookson ex Couper, Harris, p. 86. Discussion: Cookson (19476) diagnosed the sporotype Phyllocladidites without naming a type species and assigned to it the sporomorphs Phyllocladidites Mawsoni (sic) and P. Riiei (sic). In March 1953, she rejected Phyllocladidites and regarded it as a junior synonym of her new generic designate Dacrydiumites. In August 1953, Couper, apparently unaware of Cookson's action, validated Phyllocladidites and designated P. mawsonii as type species. Hence, the claim by Potonie (1958) that Phyllocladidites has priority over Dacrydiumites is correct. Cookson (1957) continued to use Dacrydiumites and included in the genus not only D. mawsonii and D. ruei but also D. florinii Cookson & Pike, 1953, D. balmei Cookson, 1957, and the forma D. mawsonii verrucosus Cookson, 1957 and further proposed in an addendum 'that all pollen grains described under the name Dacrydiumites be referred to Dacrydium . . (Cookson, 1957, p. 53). Couper's handling of Dacrydiumites in 1960 differs somewhat from the manner followed by Cookson (1957) in that he recognized D. mawsonii and D. ruei as species of Dacrydiumites but placed in the extant genus Dacrydium fossil pollen morphologically similar to Dacrydiumites florinii. Harris (1965) utilized Dacrydiumites Cookson ex Harris for

63

the reception of pollen of the D. florinii type. In fact, he designated D. florinii (not D. mawsonii) as type species, thereby making Dacrydiumites Cookson ex Harris a homonym of Dacrydiumites Cookson, which is, in turn, a synonym of Phyllocladidites Cookson ex Couper. Because Cookson in her original diagnosis emphasized the presence of localised thickenings of the exine between the proximal roots of the sacci and the body cap, species with these structures are retained in Phyllocladidites. These include P. mawsonii, P. ruei, P. reticulocassatus Harris and P. verrucosus (Cookson) Stover & Evans, comb. nov. Species previously assigned to Dacrydiumites that lack the diagnostic features (D. balmei, D. florinii and D. ellipticus Harris) are assigned to the new genus Lygistepollenites. Phyllocladidites verrucosus (Cookson) Stover & Evans, comb. nov. (Plate 2, Fig. 9) 1957 Dacrydiumites mawsonii Cookson /. verrucosus Cookson, pp. 47-48, Plate 9, Figures 15, 16. Discussion: This form is elevated to species rank because it is readily distinguished from Phyllocladidites mawsonii by having verrucated proximal polar caps. Phyllocladidites verrucosus also has a much shorter stratigraphic range than P. mawsonii, the former occurring in Palaeocene and Late Cretaceous strata in the Gippsland Basin (Tricolporites lilliei to Lygistepollenites balmei Zones). Genus Lygistepollenites Stover & Evans, gen. nov. 1965 Dacrydiumites Cookson ex Harris, p. 86. Type species: Lygistepollenites balmei (Cookson) Stover & Evans, comb, nov., here designated. Description: Pollen saccate, outline in polar view broadly elliptical, body outline circular to elliptical in polar view, elliptical in equatorial view. Grains commonly bisaccate, occasionally trisaccate; sacci pendant distally, confluent equatorially or separated; internal thickenings of sacci radial, looped, reticulate or combination thereof. Proximal cap moderately thick, scabrate to finely rugulate. Tentuitas well to poorly defined, surface psilate to granulate. Discussion: Lygistepollenites is proposed for the reception of those saccate pollen species previously assigned to Dacrydiumites by Cook-

Spec.Pubis geol.Soc.Aust., 4: pp. 55-72, Pis 1-4, 1973.

64

LEWIS E. STOVER & P. RICHARD EVANS

son (1957) which lack boss-like proximal structures and which otherwise conform to the above circumscription. The following species are included in Lygistepollenites as new combinations: L. (al. Dacrydiumites) balmei Cookson, 1957, p. 46, Plate 9, Figures 11-14; L. (al. Dacrydiumites) fiorinii Cookson & Pike, 1953, p. 479, Plate 3, Figures 20-35; and L. (al. Dacrydiumites) ellipticus Harris, 1965, p. 87, Plate 26, Figures 20, 21. Comparison: Lygistepollenites differs from Phyllocladidites Cookson ex Couper emend. Stover & Evans by lacking proximal protuberances between the body and the proximal roots of the sacci and by having generally larger sacci with coarser infrasculpture. Genus Nothofagidites Potonie, 1960 Type species: Nothofagidites flemingii (Couper) Potonie, designated by Potonie (1960, p. 132). Discussion: Except within the Tricolpites longus Zone, specimens of Nothofagidites spp. occur continuously throughout the Latrobe Group in the Gippsland Basin and are particularly abundant in the N. asperus Zone. Fossil pollen ascribed to Nothofagidites are practically indistinguishable from those of extant species of Nothofagus, except for their compressed or folded condition. Moreover, the same high degree of infraspecific and interspecific variability found within or among the pollen of living taxa is discernible in fossil specimens. Authors of Australian and New Zealand form species (Cookson 1946; Couper, 1953) have to some extent made allowances for variability in their species descriptions; however, even within the limits set by these authors, considerable difficulty arose in identifying consistently some form species and revision of some seems desirable. Other identification problems stem from the fact that each author evaluates differently the relative merit of morphologic features for identification and specific differentiation, and has couched the descriptions of these characters in different terms. Examination of Couper's preparations indicated that complete clarification of uncertainties could be achieved only through an exhaustive study involving not only the type specimens but also a large number of other specimens as well. A comprehensive study of Nothofagidites pollen is beyond the purpose and scope of this paper. We do, however, propose placing some forms in synonymy based on the examination of type specimens.

Fossil pollen of Nothofagidites are customarily arranged into groups resembling pollen of extant species informally typified by Nothofagus menziesii, N. fusca, and N. brassi (Cranwell, 1939; Cookson & Pike, 1955). Couper (1953) treated the groups essentially as subgenera without formal designation; we regard the informal groups as categories convenient for discussion and descriptive purposes. Nothofagidites pollen, Nothofagus menziesii Group Pollen of the Nothofagus menziesii Group are characterized by their relatively thin exine, deeply incised, unrimmed or weakly rimmed apertures, and comparatively large size. Two Australian form species, Nothofagidites asperus and Nothofagidites goniatus, are here included in the Nothofagus menziesii Group. Nothofagidites suggatei, included by Couper (1953, 1960) in the Nothofagus menziesii Group, has not been identified in the Gippsland Basin sections. Nothofagidites asperus (Cookson) Stover & Evans, comb. nov. (Plate 4, Fig. 6) 1959 Nothofagus asperus Cookson, p. 25, Plate 4, Figures 1, 2. Discussion: Some specimens in which the apertures are closed or only slightly opened are up to smaller in diameter than the 40//, lower size limit reported by Cookson. A majority of specimens are well within the 4060/x range. The small, low, dense grana and deep, simple apertures are diagnostic of the species. Nothofagidites goniatus (Cookson) Stover & Evans, comb. nov. (Plate 4, Fig. 4) 1959 Nothofagus goniata Cookson, p. 28, Plate 4, Figures 5, 6, Text-Fig. 5. Discussion: In her original description (as Nothofagus sp. g) Cookson (1946) stated that this species resembles closely the pollen of the Nothofagus menziesii Group in general and of Nothofagus sp. a (later validated as Nothofagus aspera) in particular. In spite of this comparison, she assigned Nothofagus sp. g to the then unnamed group that is now known as the Nothofagus brassi Group. Cookson (1959) persisted in allocating Nothofagus goniata to the Nothofagus brassi Group and

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION gave as her reason the fact that the apertures have clearly defined borders on some specimens. In our opinion, Nothofagidites goniatus shows a greater similarity to the pollen of the Nothofagus menziesii Group, especially since some of the specimens of Nothofagidites asperus have faint marginal thickenings bordering the apertures. Nothofagidites pollen, Nothofagus fusca Group Pollen of the Nothofagus fusca Group are characterized by their more or less circular equatorial outline and by the generally pronounced exinal thickening around the apertures. Two form species are included in the Nothofagus fusca Group. The one, Nothofagidites brachyspinulosus (Cookson) Harris, 1965, is known only from Australia, whereas the other, Nothofagidites flemingii (Couper) Potonie, occurs in New Zealand and Australia. Nothofagidites

flemingii (Couper) Potonie, 1960 (Plate 2, Fig. 11) 1953 Nothofagus flemingii Couper, p. 47, Plate 6, Figure 72; Plate 9, Figure 139. 1959 Nothofagus cincta Cookson, p. 26, Plate 4, Figure 3. 1960 Nothofagus flemingii Couper, Couper, p. 55, Plate 7, Figure 22. 1960 Nothofagidites flemingii (Couper) Potonie p. 132, Plate 9, Figure 196. Discussion: Couper (1953) commented on the similarity between Nothofagus flemingii and Nothofagus cincta (as N. sp. a of Cookson, 1946) and pointed out the difference in the dominant number of apertures in each species and the less deeply incised apertures in the Australian species, Nothofagus cincta. The number of apertures for Nothofagus flemingii ranged from 7-9, with 53 per cent of the specimens having 7 apertures, and 44 per cent having 8. In contrast, the number of apertures in Nothofagus cincta varied from 4-8, with a majority of specimens having 6 ox 1 apertures. Cookson (1946) presented a histogram showing the percentage of specimens versus the number of apertures for pollen of Nothofagus cincta from two localities about 900 km apart. At the Vegetable Creek locality in northeastern New South Wales, the number of apertures ranged from 5-8, with 44 per cent of the specimens having 7 apertures. At the Lai Lai locality in central Victoria, the number

65 of apertures ranged from 4-8, with 47 per cent of the specimens having 6 apertures. A comparable study using 100 specimens in a sample in which the relatively large Nothofagus fusca-type of pollen represent approximately 12 per cent of the specimens in the assemblage revealed still another variation: within the range of 4-8 apertures, 7 per cent had 4 apertures, 49 per cent had 5 apertures, 38 per cent had 6 apertures, 4 per cent had 7 apertures and only 2 per cent had 8 apertures. If Couper's main criterion for separating Nothofagidites flemingii and Nothofagus cincta is valid, then the specimens from southeastern offshore Australia represent still another species. Examination of Couper's slides in which specimens of Nothofagidites flemingii are fairly common showed that the shape, size, exine thickness, and sculpturing are essentially identical for the New Zealand and Australian specimens. From a morphologic and statistic viewpoint, and considering the variability among pollen of extant Nothofagus species, we can justify neither the retention of Nothofagidites flemingii and Nothofagus cincta as distinct taxa, nor the erection of a new species for the specimens from the Gippsland Basin. We prefer to broaden the concept of Nothofagidites flemingii with respect to aperture number (4-9), permit aperture number maxima of from 5-8, and consider Nothofagus cincta a junior synonym of Nothofagidites flemingii. With this proposal in mind, the following re-description of Nothofagidites flemingii is given.

Revised description: Pollen oblate, equatorial outline circular to subangular. Apertures equatorial, 4-9, maxima 5-8, elongate poleward, moderately incised. Exine thin, generally less than 1 fx between apertures, up to 3^ around apertures, faint indication of stratification on large specimens. Thickening of exine imparts characteristic 'U'-shaped appearance to apertures in polar view. Surface finely granulate, grana small, sparse to moderately dense, not crowded, commonly absent or widely scattered equatorially. Size, 27-54u, average diameter ca 40/JL. Comparison: Specimens of Nothofagidites flemingii are generally larger, have more delicate, smaller, more widely spaced grana, and show a greater thickening of the exine adjacent to the apertures than specimens of Nothofagidites brachyspinulosus. Occurrence: In the Gippsland Basin, the

Spec.Publs geol.Soc.Aust., 4: pp. 55-72, Pis 1-4, 1973.

LEWIS E. STOVER & P. RICHARD EVANS 66 Nothofagidites endurus Stover & Evans, species occurs consistently from the Malvacisp. nov. pollis diversus Zone, through the rest of the Latrobe Group. Scattered occurrences are (Plate 1, Fig. 7) found in the Lygistepollenites balmei Zone as Description: Pollen oblate, equatorial outwell. line subcircular to subangular. Apertures equatorial, 6-8, majority 7, colpoid, about Nothofagidites pollen, Nothofagus brassi 10^ long, moderately incised. Exine about 1/x Group thick or less between apertures, slightly thicker Pollen of the Nothofagus brassi Group are the apertures; aperture margins smooth or characterized by their relatively small size, atfinely spinulate. Surface moderately to coarsely generally subangular to angular, rarely cir- spinulate, spinules somewhat scattered, more cular equatorial outline and unthickened or less evenly distributed, equidimensional apertural margins. Among fossil fagaceous or nearly so; margins of apertures distinctly pollen in the Latrobe Group, the Nothofagus rimmed. Size 29-42^, average diameter, 36^. brassi Group is the most abundant, longest Comparison: Nothofagidites endurus differs ranging, and by far the most difficult to speciate because of the considerable varia- from N. senectus by being generally larger, bility in size, shape, aperture number, aperture having a more rounded outline, coarser sculplength, exine thickness, and the type, density, ture and thickened apertural rims. and distribution of exine sculpture among Holotype: Specimen shown on Plate 1, specimens. By using criteria set forth by Cook- Figures 7a, 7b, slide P 28087. son (1946, 1959) for the recognition and Type locality: Gippsland Basin, Esso Marlinseparation of Nothofagidites spp. we had little 2 well, Lat 38°15'59"S, Long 148°10'45" E. difficulty in identifying each species of the Type stratum: Latrobe Group at 2,340.0 m. Nothofagus brassi Group in the Proteacidites Stratigraphic range: Tricolporites lilliei into asperopolus and Nothofagidites asperus Zones. In the Malvacipollis diversus and older zones the lower part of the Malvacipollis diversus we were unable to separate certain species Zone. consistently and uniformly because of merging and overlapping characteristics. The following Nothofagidites falcatus (Cookson) Stover & species are included in the Nothofagus brassi Evans, comb. nov. Group: (Plate 4, Fig. 13) Nothofagidites deminutus (Cookson) Stover Discussion: The deeply concave margins & Evans, comb, nov.; between apertures and the sparse, relatively Nothofagidites endurus Stover & Evans, sp. large, distinct and sharply pointed spinules nov.; are diagnostic of Nothofagidites falcatus. This Nothofagidites falcatus (Cookson) Stover & species is rare in the Latrobe Group of the Evans, comb, nov.; Gippsland Basin, and is found only in the N. (al. Nothofagus) heterus Cookson, 1959, N. asperus Zone. p. 27, Plate 4, Figures 9, 10; comb, nov.; Nothofagidites senectus Dettmann & Playford, Genus Proteacidites Cookson ex Couper 1953 1968; and Type species: Proteacidites adenanthoides N. (al. Nothofagus) vansteenisii Cookson Cookson (1950), designated by Couper (1953, 1959, p. 29, Plate 4, Figure 11, comb. nov. p. 42). Nothofagidites deminutus (Cookson) Stover Proteacidites asperopolus Stover & Evans, & Evans, comb. nov. sp. nov. (Plate 4, Fig. 5) (Plate 4, Fig. 1) 1959 Nothofagus deminuta Cookson, p. 29, Plate 4, Figure 12. Description: Pollen triaperturate, heteroDiscussion: Among the N. brassi Group, this polar, one pole with prominent dome-like prospecies appears relatively late in the upper trusion, other pole unmodified or with slight part of the Malvacipollis diversus Zone and is convexity; equatorial outline triangulate to particularly abundant in the Proteacidites concavely triangular. Exine clearly to faintly asperopolus Zone, and is less common in the stratified, nexine and sexine about equally thick, each approximately 1 p in thickness; Nothofagidites asperus Zone.

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION 67 sexine incised by short, discontinuous, narrow, Nypa and described two form species from sinuous grooves that outline isolated, irregu- Sarawak, of which he selected S. echinatus larly shaped patches of sexinous material, par- as type species. Spinizonocolpites echinatus is ticularly in the non-apertural areas; apertural interpreted as conspecific with the taxa identiareas finely reticulate. Columella moderately fied and described as Monosulcites prominadense and distinct around apertures, generally tus Mclntyre (1965) and as Baltisphaeridium less dense on other parts of the pollen. Aper- taylori Cookson & Eisenack (1965Z>). tures equatorial, porate, pores about 1.5^ Genus Tricolpites Cookson ex Couper, 1953 across in polar view. Equatorial diameter, 30Type species: Tricolpites reticulatus Cook42jn, mean 3 8 s e v e r a l hundreds of specison, designated by Couper (1953, p. 61). mens observed. Discussion: The more coarsely sculptured longus Stover & Evans, sp. nov. parts of the sexine that correspond also to Tricolpites (Plate 1, Figs 13, 14) those areas having relatively few columellae Description: Pollen triaperturate, brevaxial, tend to become detached. Consequently, specimens with the sexine intact only around the equatorial outline triangular with straight to apertures or in other areas of fine sculpture gently concave or convex sides, apertural corners narrowly rounded. Exine about 1.5^ are common in some preparations. not clearly stratified and apparently Comparison: Proteacidites asperopolus is thick, surface smooth to faintly similar to P. pachypolus Cookson & Pike, 1954 lacks columellae; or roughened and usually with triin size, equatorial outline, and in being strongly scabrate radiate scar on one, more rarely, on both heteropolar. Although both species are finely surfaces. Apertures orequatorial, colpi short, reticulate in the apertural areas, the sexine is narrow, sharply defined with apertural margins uniformly reticulate over the entire surface incurved. Size, 39-50/z, average, 4 3 c a 30 on specimens of P. pachypolus (see Plate 3, specimens. Fig. 8), whereas on specimens of P. asperoComparison: Tricolpites longus differs from polus the sculpture is much coarser and nonreticulate in the polar and interapertural areas. T. sabulosus Dettmann & Playford, 1968 by being larger and by having a smooth to scabHolotype: Specimen shown on Plate 4, rate exine rather than coarse and fine grana Figures la, lb, slide P 28049. distributed uniformly over the surface. Type locality: Bass Basin, Esso Bass-2 well, Types: Holotype, Plate 1, Figures 14a, 14b, Lat 39° 53' 09" S, Long 146° 18' 18" E. slide P 28099; paratype, Plate 1, Figure 13, Type strata: Eastern View Coal Measures slide P 28100, both specimens from the same at 1,313.9 m. locality and sample. Type locality: Gippsland Basin, Esso Genus Spinizonocolpites Muller, 1968 Flounder-2 well, Lat 38°19'18"S, Long Type species: Spinizonocolpites prominatus 148°23'53"E. (Mclntyre), Stover & Evans, comb, nov., = Type stratum: Latrobe Group at 2,531.0 m. S. echinatus Muller and designated by Muller Stratigraphic range: Tricolporites lilliei and (1968, p. 11) as type species. Tricolpites longus Zones. Spinizonocolpites prominatus (Mclntyre) Genus Tricolporites Cookson 1947 Stover & Evans, comb. nov. Type species: Tricolporites sphaerica Cook(Plate 3, Fig. 5) monotypic when proposed (Cookson 1965 Monosulcites prominatus Mclntyre, p. son, 1947a, p. 195) 214, Figures 33, 34. 1965 Baltisphaeridium taylori Cookson & Tricolporites lilliei (Couper) Stover & Evans, Eisenack, p. 137, Plate 16, Figures 9-11. comb. nov. (Plate 1, Fig. 8) 1968 Spinizonocolpites echinatus Muller, pp. 11, 12, Plate 3, Figure 3. 1953 Tricolpites lilliei Couper, p. 62, Plate Discussion: Muller (1968) proposed Spini8, Figures 116, 117. zonocolpites for the reception of fossil pollen 1960 Tricolpites lilliei Couper, Couper, p. 64, similar to the pollen of the extant palm genus Plate 10, Figure 19. SpecJPubls geoLSoc.Aust, 4: pp. 55-72, Pis 1-4, 1973.

68

LEWIS E. STOVER & P. RICHARD EVANS Discussion:

Specimens of Tricolporites

lilliei

in the Australian Late Cretaceous and Early to Middle Palaeocene show considerable variability in colpi development, and in the density of the spines, and in the clarity of the internal aperture. The colpi on some specimens are relatively short, whereas on others they extend

far into the polar areas; however, splitting of the exine at the ends of the colpi is fairly common, so that the apertures may appear longer than they really are. The ora are only faintly developed and are at best displayed on specimens that are not compressed parallel to the equatorial plane. Spine development is

TABLE I I

Alphabetical listing of illustrated specimens Name Aglaoreidia sp. A Australopollis obscurus Beaupreaidites elegansiformis Camarozonosporites ohaiensis Camarozonosporites sp. A Caryophyllidites polyoratus Cupanieidites orthoteichus Ischyosporites sp. A Kuylisporites waterbolkii Liliacidites sp. A Lygistepollenites balmei Malvacipollis diversus Monosulcites sp. A Myrtaceidites tenuis Nothofagidites asperus Nothofagidites deminutus Nothofagidites endurus Nothofagidites falcatus Nothofagidites flemingii Nothofagidites goniatus Nothofagidites senectus Ornamentifera sentosa Peromonolites densus Phyllocladidites verrucosus Planar tetrad Polycolpites sp. A Proteacidites annularis Proteacidites asperopolus Proteacidites dilwynensis Proteacidites grandis Proteacidites pachypolus Proteacidites palisadus Proteacidites sp. A Proteacidites sp. B Rugulatisporites sp. A Santalumidites cainozoicus Spinizonocolpites prominatus Stereisporites regium S. (Tripunctisporis) sp. Tetracolporites sp. A Tetrahedral tetrad Tiliaepollenites notabilis Tricolpites gillii Tricolpites longus Tricolpites waiparaensis Tricolpites sp. A Tricolpites sp. B Tricolporites lilliei Tricolporites sp. A Triorites edwardsii Triorites harrisii Triorites magnificus Triporopollenites sp. A

Locality

Depth

Slide No.

G roper-1 Tuna-3 Bass-2 Barracouta-1 Wahoo-1 Bass-2 Groper-1 Bass-1 Bass-3 Bass-2 Alberton W-138 Bass-2 Bass-2 Marlin-3 Bass-3 Bass-2 Marlin-2 Groper-1 Cod-1 Bass-2 Tuna-3 Flounder-2 Marlin-2 Snapper-2 Wahoo-1 Bass-2 Bass-2 Bass-2 Marlin-1 Bass-2 Marlin-3 Wahoo-1 Barracouta-1 Bass-2 Bass-2 Bass-2 Tasmania* Bass-2 Tuna-3 Snapper-2 Bass-2 Marlin-3 Bass-2 Flounder-2 Wahoo-1 Tuna-1 Bass-2 Barracouta-1 Bass-2 Wahoo-1 Bass-2 Victoria* Barracouta-1

940-3 m 1,518-2 m 1,417-6 m 2,653-5 m 576-5 m 1,668-0 m 940-3 m 1,641-6 m 1,377-4 m 1,218-8 m 61-6 m 1,417-6 m 1,313-9 m 1,563-7 m 1,527-7 m 930-3 m 2,340-0 m 966-9 m 1,948-0 m 1,472-6 m 2,028-8 m 2,531 Om 2,257-6 m 2,001-1 m 562-4 m 1,505-8 m 1,313-9 m 1,313-9 m 1,418-8 m 1,313-9 m 1,551-8 m 576-5 m 2,211-6 m 1,263-3 m 1,417-6 m 1,155-2 m

P 28109 P 28115 P 28111 P 28060 P 28103 P 28049 P 28109 P 28057 P 28059 P 28112 P 28142 P 28050 P 28049 P 28074 P 28096 P 28086 P 28087 P 28141 P 29069 P 28056 P 28120 P 29099 P 28048 P 28116 P 28101 P 28055 P 28049 P 28049 P 28067 P 28049 P 29076 P 28104 P 28091 P 28073 P 28050 P 28070 P 28114 P 28052 P 28117 P 28118 P 28049 P 28084 P 28112 P 28099, 100 P 28121 P 28101 P 28066 P 28060 P 29090 P 28104 P 28049 P 28113 P 28093

—

1,592-0 m 1,954-7 m 2,499-2 m 1,313-9 m 1,569-5 m 1,417-6 m 2,531-0 m 576-5 m 1,807-7 m 1,218-8 m 2,653-5 m 1,394-8 m 567-5 m 1,313-9 m —

2,652-0 m

* Surface sample from locality given in Cookson & Eisenack (1965a).

Plate

Figure

4 1 3 1 2 1 3 3 3 4 2 3 4 3 4 4 1 4 2 4 1 2 2 2 2 2 2 4 3 3 3 1 2 3 3 4 3 1 2 2 4 3 1 1 2 2 4 1 4 1 2 4 1

11 3 6 1 5 5 2 14 13 2 8 3 9 1 6 5 7 13 11 4 4 1 7 9 4 10 13 1 9 10 8 9, 10 14 7 11,12 3 5 11 12 3 7 4 6 13, 14 2 6 10 8 8 2 15 12 12

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION 69 erratic; some individuals are nearly devoid of of the palynology of the La Trobe No. 1 bore, spines, and others have numerous but rather Victoria. Permission to publish the information widely scattered spines. Because of the col- derived from offshore Gippsland Basin wells porate nature of the apertures, Tricolpites has been granted by Esso Australia Ltd* and lilliei is transferred to Tricolporites. Broken Hill Proprietary Co. Ltd, for which we are most appreciative. Mr A. D. Partridge read ACKNOWLEDGMENTS the manuscript and offered helpful suggestions. In the course of this study, we have been aided and encouraged by a number of individuals, both in Australia and in the U.S.A., ILLUSTRATED SPECIMENS to whom we are sincerely grateful. During Specimens illustrated on Plates 1-4 are listed the early stages Dr Mary E. Dettmann (Uni- in Table II, together with the well name and versity of Queensland) and Mr W. K. Harris depth from which each specimen was obtained. (Department of Mines, South Australia) were These are in the palaeontologic collections most helpful in exchanging ideas concerning in the National Museum of Victoria, Melthe identification of many problematical taxa bourne, and the slides have been assigned as well as new forms. Mr J. S. Bain (Esso accession 'P' numbers. The exact locations of Standard Oil (Australia) Ltd) provided an the specimens are on file with the slides. Beexcellent suite of Palaeocene and Eocene sur- cause generally better preserved specimens face samples from southwestern Victoria which were recovered from some of the Bass rather yielded much needed comparative material, than Gippsland Basin wells, we have comand Miss Joan B. Stough (Exxon Company, monly chosen the more favourable specimens U.S.A.) assisted materially through her study from the Bass Basin for illustration. REFERENCES BERGGREN, W. A., 1969: Rates of evolution in , 19656: Microplankton from the Paleocene

some Cenozoic planktonic Foraminifera. Pebble Point Formation, south-western VicMicropaleontology, 15, pp. 351-365. toria, Pt 1. Proc. R. Soc. Vict., 78, pp. 137141. COOKSON, I. C., 1946: Pollen of Nothofagus Blume from Tertiary deposits in Australia. COOKSON, I. C., & EISENACK, A., 1958: Microplankton from Australian and New Guinea Proc. Linn. Soc. N.S.W., 77, pp. 49-63. Upper Mesozoic sediments. Proc. R. Soc. , 1947a: On fossil leaves (Oleaceae) and Vict., 70, pp. 19-79. a new type of fossil pollen grain from Aus, , 1965a: Microplankton from the tralian brown coal deposits. Proc. Linn. Soc. Browns Creek Clays, SW. Victoria. Proc. R. N.S.W., 72, pp. 183-197. Soc. Vict., 79, pp. 119-131. , 1947&: Plant microfossils from the , , 19656: Microplankton from the lignites of Kerguelen Archipelago. Dartmoor Formation, SW. Victoria. Proc. R. Br.Aust.N.Z. Antarctic Res. Exped., 1929Soc. Vict., 79, pp. 133-137. 1931, Reps., ser. A, 2, (8), pp. 129-142. , , 1965c: Microplankton from the , 1950: Fossil pollen grains of proteaceous Paleocene Pebble Point Formation, southtype from Tertiary deposits in Australia. western Victoria, Pt 2. Proc R. Soc. Vict., Aust. J. Res., ser. B, 3, pp. 166-176. 79, pp. 139-146. , , 1967A: Some Early Tertiary , 1953: The identification of the sporomicroplankton and pollen grains from a morph Phyllocladidites with Dacrydium and deposit near Strahan, western Tasmania. its distribution in southern Tertiary deposits. Proc. R. Soc. Vict., 80, pp. 131-140. Aust. J. Bot., 7, pp. 64-70. -, 1967b: Some microplankton , 1957: On some Australian Tertiary spores from the Paleocene Rivernook Bed. Proc. and pollen that extend the geological and R. Soc. Vict., 80, pp. 247-257. geographical distribution of living genera. COOKSON, I. C., & PIKE, K . M., 1953: A contriProc. R. Soc. Vict., 69, pp. 41-53. bution to the Tertiary occurrence of the , 1959: Fossil pollen grains of Nothofagus genus Dacrydium in the Australian region. from Australia. Proc. R. Soc. Vict., 71, pp. Aust. J. Bot., 1, pp. 474-484. 25-30. , , 1954: Some dicotyledonous , 1965a: Cretaceous and Tertiary micropollen types from Cainozoic deposits in the plankton from southeastern Australia. Proc. Australian region. Aust. J. Bot., 2, pp. 197R. Soc. Vict., 78, pp. 85-93. 219. * Esso Australia Ltd also contributed generously to the cost of publication. Spec.Publs geoI.Soc.Aust., 4: pp. 55-72, Pis 1-4, 1973.

LEWIS E. STOVER & P. RICHARD EVANS

70

? ? 1955: The pollen morphology of Nothofagus Bl. subsection Bipartitae Steen. Aust. J. Bot., 3, pp. 197-206. COUPER, R. A., 1953: Upper Mesozoic and Cainozoic spores and pollen grains from New Zealand. Paleont. Bull, Wellington, 22, pp. 1-77. , 1960: New Zealand Mesozoic and Cainozoic plant microfossils. Paleont. Bull., Wellington, 32, pp. 1-87. CRANWELL, L. M., 1939: Southern beech pollen. Rec. Auckland Inst. Mus., 2, pp. 175-196. DAVEY, R. J., et al1966: Studies on Mesozoic and Cainozoic dinoflagellate cysts. Bull. Br. Mus. nat. Hist. Geol., supp. 3, pp. 1-248.

DEFLANDRE, G . , & COOKSON, I. C . , 1 9 5 5 :

Fossil

microplankton from Australian Late Mesozoic and Tertiary sediments. Aust. J. mar. Freshwat. Res., 6, pp. 242-313. DETTMANN, M . E . , & PLAYFORD, G . , 1 9 6 8 : T a x o -

nomy of some Cretaceous spores and pollen grains from eastern Australia. Proc. R. Soc. Vict., 81, pp. 69-94. , 1969: Palynology of the Australian Cretaceous—a review: in Campbell, K. S. W. (Ed.), Stratigraphy and Palaeontology: Essays in Honour of Dorothy Hill. A.N.U. Press, Canberra. DORING, H., et al., 1966: X)ber einige neue Subformgenera der Sporengattung Stereisporites Th. & Pf. aus dem Mesozoikums and Alttertiar Mitteleuropes. Geologie, 55, pp. 72-89. DRUGG, W. S., 1967: Palynology of the Upper Moreno Formation (Late Cretaceous-Paleocene), Escarpado Canyon, California. Palaeontographica B, 120, pp. 1-71. HARRIS, W. K., 1965: Basal Tertiary microfloras from the Princetown area, Victoria, Australia. Palaeontographica B, 115, pp. 76-106. 1966: Proteacidites latrobensis. Taxon, 15, pp. 332-333. JAMES, E . A . , & EVANS, P . R . , 1 9 7 1 : T h e

strati-

graphy of the offshore Gippsland Basin. J. Aust. Petrol. Expl. Assn, 77, pp. 71-74. JENKINS, D. G., 1971: New Zealand Cenozoic planktonic foraminifera. Palaeont. Bull., Wellington, 42, pp. 1-278. KRUTZSCH, W., 1959: Zur Kenntnis der praquartaren periporaten Pollenformens. Geologie, 75, pp. 16-72.

Lewis E. Stover*, P. Richard Evans, Esso Australia Ltd., G.P.O. Box 4047, Sydney, New South Wales 2001.

LUNT,

C.

K.,

&

JAMES,

E.

A.,

1968:

Esso

Nautilus-1 well completion report. Rep. Aust. Petrol. Search Subsidy Acts, [unpublished]. MCGOWRAN, B., 1965: Two Paleocene foraminiferal faunas from the Wangerrip Group, Pebble Point coastal section, western Victoria. Proc. R. Soc. Vict., 79, pp. 9-74. MCINTYRE, D. J., 1965: Some new pollen species from New Zealand Tertiary deposits. N.Z.JI Bot.,

3, p p .

204-215.

MULLER, J., 1968: Palynology of the Pedawan and Plateau Sandstone Formations (Cretceous-Eocene) in Sarawak, Malaysia. Micropaleontology, 14, pp. 1-37. POTONIE, R., 1956: Synopsis der Gattungen der Sporae dispersae, I Teil. Geol. Jb., Beih., 23, pp.

1-103.

, 1958: Synopsis der Gattungen der Sporae dispersae, II Teil. Geol. Jb., Beih., 31, pp. 1-144.

, 1960: Synopsis der Gattungen der Sporae dispersae, III Teil. Geol. Jb., Beih., 39, pp. 1-189. RICHARDS, K . A . , & HOPKINS, B. M . ,

1969:

Ex-

ploration in the Gippsland, Bass and Otway Basins, Australia. ECAFE. Nat. Resour. Symp. Dev. Petrol. Resour. Asia Far East, Canberra, [unpublished] STOVER, L. E., 1973: Palaeocene and Eocene species of Deflandrea (Dinophyceae) in Victorian coastal and offshore basins, Australia. Spec. Pubis geol. Soc. Aust., 4, pp. 167-188. TAYLOR, D. J., 1966: The mid-Tertiary foraminiferal sequence; in Esso Gippsland Shelf No. 1. Bur. Miner. Resour., Aust. Petrol. Search Subsidy Acts. Publ., 76, pp. 31-44. TE PUNGA, M. J., 1948: Nothofagus pollen from the Cretaceous Coal Measures at Kaitangata, Otago, New Zealand. N.Z.JI Sci. Tech., 29B, pp. 32-35.

WALLIS, W. E., 1967: Offshore petroleum exploration, Gippsland and Bass Basins, southeast Asia. Proc. 7th World Petrol. Congr., 2, pp.

783-791.

WEEKS, L . G . , & HOPKINS, B. M . , 1 9 6 6 :

Geology

and exploration of three Bass Strait basins. Bull. Am. Ass. Petrol. Geol., 51, pp. 742-760. WILSON, G. J., 1967: Some species of Wetzeliella Eisenack (Dinophyceae) from New Zealand Eocene and Paleocene strata. N.Z.JI Bot., 5, pp. 469-497.

* Present address: Esso Production Research Co., P>0. Box 2189, Houston, Texas 77001, U.S.A.

UPPER CRETACEOUS-EOCENE SPORE-POLLEN ZONATION EXPLANATION OF PLATES PLATE 1

Typical spores and pollen of the Nothofagidites senectus (Figs 1-6) and Tricolporites lilliei Zones Fig. 1. Camarozonosporites ohaiensis (Couper) Dettmann & Playford, 1968, interference contrast, X 400. Fig. 2. Thorites edwardsii Cookson & Pike, 1954 interference contrast, X 1000. Fig. 3. Australopollis obscurus (Harris) Krutzsch, 1966, interference contrast, X 800. Fig. 4. Nothofagidites senectus Dettmann & Playford, 1968, specimen shown at two focus levels, bright field, X 800. Fig. 5. Caryophyllidites polyoratus Couper, 1960, interference contrast, X 800. Fig. 6. Tricolpites gillii Cookson, 1957, interference contrast, X 800. Fig. 7. Nothofagidites endurus Stover & Evans, sp. nov., holotype at two focus levels, interference contrast, X 800. Fig. 8. Tricolporites lilliei (Couper) Stover & Evans, comb, nov., interference contrast, X 800. Figs 9, 10. Proteacidites palisadus Couper, 1953. 9, specimen at intermediate focus level; 10, specimen at high (10a) and intermediate (10b) focus levels; both specimens in bright field and at X 800. Fig. 11. Stereisporites regium (Drozh.) Drugg, 1967, at proximal (11a) and distal (lib) focus levels, interference contrast, X 800. Fig. 12. Triporopollenites sp. A, bright field, X 800. Figs 13,14. Tricolpites longus Stover & Evans, sp. nov., 13, paratype, bright field, X 800; 14, holotype at intermediate (14a) and high (14b) focus levels, interference contrast, X 800. PLATE 2

Typical spores and pollen of the Tricolpites longus (Figs 1-6) and Lygistepollenites balmei Zones Fig. 1. Ornamentifera sentosa Dettmann & Playford, 1968, bright field, X 500. Fig. 2. Tricolpites waiparaensis Couper, 1960, specimen at high (2a) and intermediate (2b) focus levels in bright field, X 800. Fig. 3. Tetracolporites sp. A, interference contrast, X 800. Fig. 4. Planar tetrad, bright field, X 800. Fig. 5. Camarozonosporites sp. A, distal (5a) and proximal (5b) surfaces in interference contrast, X 400. Fig. 6. Tricolpites sp. A, interference contrast, X 1000. Fig. 7. Peromonolites densus Harris, 1965, at intermediate (7a) and low (7b) focus levels bright field, X 800. Fig. 8. Lygistepollenites balmei (Cookson) Stover & Evans, comb, nov., hypotype in interference contrast, X 325. Fig. 9. Phyllocladidites verrucosus (Cookson) Stover & Evans, comb, nov., hypotype in interference contrast, X 800. Fig. 10. Polycolpites sp. A, interference contrast, X 500. Fig. 11. Nothofagidites flemingii (Couper) Potonie, 1960, at high (11a) and intermediate (lib) focus levels in interference contrast, X 800. Fig. 12. Stereisporites (Tripunctisporis) sp. in bright field, X 800. Fig. 13. Proteacidites annularis Cookson, 1950, interference contrast, X 800. Fig. 14. Proteacidites sp. A, interference contrast, X 800. Fig. 15. Thorites harrisii Couper, 1953, interference contrast, X 800. Spec.Publs geol.Soc.Aust., 4: pp. 55 72, Pis 1-4, 1973.

72

LEWIS E. STOVER & P. RICHARD EVANS PLATE 3

Typical spores and pollen of the Malvacipollis diversus Zone Fig. 1. Myrtaceidites tenuis Harris, 1965, interference contrast, X 800. Fig. 2. Cupanieidites orthoteichus Cookson & Pike, 1954 at intermediate (2a) and high (2b) focus levels in bright field, X 800. Fig. 3. Malvacipollis diversus Harris, 1965, bright field, X 800. Fig. 4. Tiliaepollenites notabilis Harris, 1965, interference contrast, X 500. Fig. 5. Spinizonocolpites prominatus (Mclntyre) Stover & Evans, comb, nov., view of apertural face (5a) and of exinal surface (5b), interference contrast, X 800. Fig. 6. Beaupreaidites elegansiformis Cookson, 1950, interference contrast, X 800. Fig. 7. Proteacidites sp. B at high (7a) and intermediate (7b) focus levels, bright field, X 800. Fig. 8. Proteacidites pachypolus Cookson & Pike, 1954, interference contrast, X 800. Fig. 9. Proteacidites dilwynensis Harris, 1965, interference contrast, X 640. Fig. 10. Protcacidites grandis Cookson, 1950, bright field, X 640. Figs 11, 12. Rugulatisporites sp. A, proximal (11) and distal (12) surfaces in interference contrast, X 500. Fig. 13. Kuylisporites waterbolkii Potonie, 1956, interference contrast, X 800. Fig. 14. lschyosporites sp. A, interference contrast, X 800. PLATE 4

Typical spores and pollen of the Proteacidites asperopolus (Figs 1-10) and Nothofagidites asperus Zones Fig. 1. Proteacidites asperopolus Stover & Evans, sp. nov., holotype at intermediate (la) and low (lb) focus levels, interference contrast, X 800. Fig. 2. Liliacidites sp. A, bright field, X 800. Fig. 3. Santalumidites cainozoicus Cookson & Pike, 1954, interference contrast, X 800. Fig. 4. Nothofagidites goniatus (Cookson) Stover & Evans, comb, nov., hypotype, interference contrast, X 800. Fig. 5. Nothofagidites deminutus (Cookson) Stover & Evans, comb, nov., hypotype, interference contrast, X 800. Fig. 6. Nothofagidites asperus (Cookson) Stover & Evans, comb, nov., hypotype, interference contrast, X 800. Fig. 7. Tetrahedral tetrad, interference contrast, X 800. Fig. 8. Tricolporites sp. A, interference contrast, X 800. Fig. 9. Monosulcites sp. A, (Dyad), interference contrast, X 800. Fig. 10. Tricolpites sp. B, interference contrast, X 800. Fig. 11. Aglaoreidia sp. A, interference contrast, X 800. Fig. 12. Triorites magnificus Cookson, 1950, bright field, X 800. Fig. 13. Nothofagidites falcatus (Cookson) Stover & Evans, comb, nov., hypotype at high (13a) and intermediate (13b) focus levels in bright field, X 800.

LEWIS E . STOVER & P . RICHARD EVANS

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE 1

PLATE

2

L E W I S E . STOVER & P . RICHARD EVANS

L E W I S E . STOVER & P . RICHARD EVANS

Spec.Publs geol.Soc.Aust., 4, 1973,

PLATE 3

PLATE

4

L E W I S E . STOVER & P . RICHARD EVANS

REAPPRAISAL OF SOME PALYNOMORPHS OF SUPPOSED PROTEACEOUS AFFINITY. I. THE GENUS BEA UPREAIDITES COOKSON EX COUPER AND THE SPECIES PROTEACIDITES HAKEOIDES COUPER By A. R. H . M A R T I N

(With 1 Plate and 2 Text-Figures) ABSTRACT

The Tertiary palynomorph genus Beaupreaidites Cookson ex Couper, 1953 is provided with an emended diagnosis. The Tertiary palynomorph species Proteacidites hakeoides Couper, 1960, is reinterpreted and provided with an emended diagnosis. This form is shown to be unlike the extant genus Hakea Schrad. with which it had been compared. The definition of palynomorphs in terms of supposed taxonomic affinity is criticised. The phytogeographical implications of fossil pollen grains resembling those of the Proteaceae are briefly discussed. INTRODUCTION

The palynomorph genera Beaupreaidites and Proteacidites were proposed by Cookson (1950) for the reception of fossil triaperturate pollen grains apparently referable to the family Proteaceae. The earlier species described were Australian or New Zealand forms and could, without raising disturbing questions, be assumed to be of actual proteaceous affinity. In recent years both genera were widely reported in the northern hemisphere far outside the present geographical range of Proteaceae, and in sediments of Senonian and Maestrichtian age (Samoilovich, 1967; Srivastava, 1969), whereas the southern hemisphere forms were mostly Palaeocene or younger. Complex issues of migration have been raised but more important is the question of whether the grains are acceptable evidence of the Proteaceae. The pollen morphology of modern Proteaceae is not strictly uniform nor is it unique. Pollen forms as different as those of Protea (Plate 1, Fig. 1), Grevillea (Plate 1, Fig. 3), Carnarvonia (Plate 1, Fig. 4), Telopea (Plate 1, Fig. 5), Aulax (Plate 1, Fig. 6) and Franklandia (Plate 1, Fig. 8) are all included in the family and it becomes obvious that if a genus such as Proteacidites is to be treated as a morphological entity, it is very difficult if not impossible to define it in terms which will exclude other triporate palynomorphs; while if ascription to the genus is to depend on 'affinity', however that is to be estabished, then the genus cannot be morphologically homogeneous. The intention implicit in Cookson's (1950) account and explicitly stated by Couper (1953) was presumably to exclude Spec.Publs geol.Soc.Aust., 4: pp. 73-78, PL 1, 1973.

from Proteacidites those forms distinctive enough to place in other proteaceous genera. This course was followed in defining the genera Banksieaeidites Cookson, 1950, and Beaupreaidites Cookson ex Couper, 1953. Couper (1953) recognised Knightia R.Br., an extant genus, alongside Proteacidites, though it is hard to see any point in which the fossil Knightia pollen is excluded morphologically from Proteacidites as defined. Other forms are given specific names only suggestive of generic affinity, e.g. P. hakeoides Couper, 1960. Very clearly, no consistency has been observed in the application of nomenclature. A considerable number of putatively proteaceous palynomorphs have now been placed in the two genera Proteacidites and Beaupreaidites, and on this basis, long since discredited for leaf fossils, wide implications as to the former distribution of Proteaceae have been drawn. In this study and a succeeding one a preliminary attempt will be made to assess the validity of certain 'proteaceous' palynomorphs as indicators of the occurrence of Proteaceae, and to establish more homogeneous taxonomic groupings.

THE GENUS

BEAUPREAIDITES

Unlike Proteacidites, which from the outset comprised a very miscellaneous group, Beaupreaidites, when initially proposed, was a clearly circumscribed, albeit untypified, morphological category (Cookson, 1950). Couper (1953) validated the genus, designating B. elegansiformis Cookson as the type, and did not describe any further species. Cookson's

74

A. R. H. MARTIN

diagnosis was not explicit but the intention was clearly to form a receptacle for pollen grains with a close morphological resemblance to the pollen of the extant genus Beauprea Br. & Gr. ('. . . it shows a remarkable resemblance to the pollen of two species of Beauprea\ B. elegansiformis was stated to be 'closely similar in all respects to pollen of Beauprea elegans'—Cookson, 1950). These similarities are borne out by the figures accompanying the paper. The genus was subsequently reported from New Zealand, U.S.S.R., and the British Isles, but without overt extension of its morphological bounds. Srivastava (1969) emended the genus to include certain species described by U.S.S.R. palynologists as species of Proteacidites. As these forms were colporate or colporoidate (Fig. la) and Proteacidites is by definition porate, removal from the latter genus was fully correct. Tschudy (1971) considers the emendation invalid as the type species Beaupreaidites elegansiformis Cookson lacks vestibulate apertures, and would be excluded from the genus. This is in fact the case; the emendation was intended to include only certain additional types of surface ornamentation and the vestibulate character may have been taken for granted, as neither the original description of Beaupreaidites nor the somewhat revised description given by Potonie (1960) specified that vestibules were absent. It is clear, however, from photographs of the type (Cookson, 1950, Plate 1, Fig. 4) and from the modern genus Beauprea (Plate 1, Fig. 11), which, from the wording of the

description has to be taken into consideration, that this palynomorph is not vestibulate. Nor is it colpate, pace Tschudy (1971), but colpoidate. In order to bring about a better understanding of the morphology of Beaupreaidites, the following emended diagnosis is submitted. Genus Beaupreaidites Cookson ex Couper, 1953, emend. (Cookson, 1950, Plate 1, Fig. 2) Emended diagnosis: Pollen grains triaperturate, subangular to semiangular in polar view, oblate or nearly so in equatorial view, more or less isopolar; apertures forming short colpoids, meridionally elongated, with irregular thin margins and tapering exine, angles smoothly rounded; internally the aperture bounded by an area or zone ('solution', Thomson & Pflug, 1953) of roughened, foveolate or warty endexine, corresponding to the postatrium of a porate grain; annulus, vestibule and costae absent; exine reticulate, foveolate or areolate, often the reticulum combined with raised low verrucae; size of grain medium to rather large. Remarks'. The size of the grains as given by Cookson is 38-52/i. for B. elegansiformis and is not stated for B. verrucosus. The distinctive character of the colpoid found both in Beaupreaidites and in the pollen of Beauprea is shown in Figure 1A and Plate 1, Figure 11. The reconstruction of the colpoid of Beaupreaidites Cookson ex Couper emend.

Fig. 1 (A) Construction of aperture of Beauprea and of Beaupreaidites, showing combination of colpoid and post-atrium. (B) Construction of aperture of Beaupreaidites sensu Srivastava ( Siberiapollis Tschudy in part), showing its colporate character. After Srivastava, redrawn, co = colpus; os = os.

REAPPRAISAL OF SOME PALYNOMORPHS 75 Srivastava is as shown in Figure IB. The two resemblances to some modern families may be do not correspond. Beaupreaidites elegansi- seen, it is unlikely that any modern family is formis sensu Samoilovich in Samoilovich et al., represented. They bear no resemblance in their 1961, does not correspond with Australian apertural structure to Proteaceae, with which material given this name by Cookson. The they have hitherto been compared. Some Sapinplacing of several of the formerly described daceae {e.g. Pometia, Plate 1, Fig. 9), species of Proteacidites from western Siberia Eleagnaceae and Symplocaceae have apertures in the new palynomorph genus Siberiapollis, by more or less similar to Siberiapollis or to the Tschudy (1971), leaves open the question of forms listed above. whether certain other North American or Siberian species may be correctly assigned to THE SPECIES PROTEACIDITES HAKEOIDES either Proteacidites or Beaupreaidites. From an COUPER examination of the description and photoThe palynomorph species Proteacidites graphs in Samoilovich et al. (1961) and Srivas- hakeoides was described by Couper (1960) tava (1969) I am of the opinion that the from Eocene of New Zealand. It following species cannot be correctly assigned was said to besediments with the pollen to either genus: Proteacidites convexiporus grains of some 'comparable recent species of Hakea in Samoilovich et al., P. constrictus Samoilovich shape, nature of aperture and sculpture'. The et al., P. mollis Samoilovich et al., P. bellus occurrence of a member of the Proteaceae Samoilovich et al, P. crispus Samoilovich et allied to the, now purely Australian, genus al., P. formosus Samoilovich et al. and Beau- Hakea in New Zealand as early as the Early preaidites libitis Srivastava. Some of these are Eocene would be geographically very interperhaps assignable to Siberiapollis, others will esting since the two areas are thought to have require new generic standing, but all are of had only slight geographical contact at this more complex pore structure than any puta- period and by a northern route. Evidences of tively proteaceous genus. floristic migration attributable to such contact Affinity: These grains are all in the Norma- are not numerous and some are inferential polles, an artificial grouping of palynomorph rather than concrete palaeobotanical records. From examination of the type slide of this genera characterised by complex (often 'bizarre', sensu Thomson & Pflug, 1953), often species, bearing two specimens, one the holovestibulate or polyvestibulate apertures. While type, it appeared that the structure of the

Fig. 2. Construction of Proteacidites hakeoides: drawn from holotype. Spcc.Publs geal.Soc.Aust., 4: pp. 73-78, PI. 1, 1973.

76

A. R. H. MARTIN

aperture was very different from that of Hakea. As this feature has been only briefly described by Couper, an emended description of the species based on these two grains, is given here. Proteacidites hakeoides Couper, 1960 descr. emend. (Fig. 2) Emended diagnosis'. Exine verrucose, verrucae ca 1.0-1.5^ high and up to 2jx diam., irregular, rarely approximately circular in outline, arising from thin tectum which is hard to discern owing to overlapping of verrucae lying outside the plane of focus; each verruca supported on ca 2-5 narrow columellae 1 ^ in length; total thicknes of exine including verrucae ca 4.5-5.0/x in interporia and ca 5.5/* at lower rim of pore aspis; pore region consisting of domed aspides, devoid of verrucae, but appearing to have very thin ectexine, perhaps only ca 0.2^ thick, supported on minute columellae which decrease in size towards apex of dome; apex containing a pore aperture ca 5.5-7.5//, in diam., at margins of which remains of thin membrane may be detectable (Fig. 2); interior of aspis ca 10-11/* in diam. Remarks: The reconstruction of the sectional view may not be strictly accurate as it indicates as sharply focussed, verrucae which, in fact, lie slightly below or above the plane. It does give a reasonable depiction of the visual impression obtained however, as the eye adjusts to seeing these slightly unfocussed features. Thus the apertures of Proteacidites hakeoides differ from those of Hakea (see for example, Erdtman, 1966, Fig. 210) in that the pore is a clearly defined region in the apex of the domed aspis, which is a structural continuation of the complete exine. In Hakea, the pore aperture appears always to be defined by the basal limit of the domed expanded pore membrane, which, though it may be of considerable thickness, is not directly comparable with the aspis of P. hakeoides, and is normally broken off as a whole when the grain germinates. The remnants of thin pore membrane show that in P. hakeoides a germ tube grew out through the restricted apical aperture of the aspis. The dimensions of the ora given by Couper correspond most closely with the internal diameter of the aspis which suggests that the somewhat inconspicuous true apertures had not been seen. The statement that the

sculpture is comparable with that of the pollen grains of some Hakea species also needs qualification. Hakea marginata and H. ruscifolia (illustrated in Erdtman, 1966, Figs 212 a & b) differ in having less closely clustered sculptural elements. Other species of Hakea may have subtectal columellae comparable in diameter and distribution over the exine with the verrucae of P. hakeoides, but these species lack supratectal sculpture. A close resemblance to the structure of the aperture of P. hakeoides is seen in that group of Proteacidites species which includes P. adenanthoides, P. crassus, P. grandis, and an, apparently unnamed, species seen in Cookson's Comaum material, perhaps allied to P. obscurus (Plate 1, Fig. 10). There is thus no reason to suppose that a grevilleoid genus allied to Hakea ever grew in New Zealand.

DISCUSSION The great majority of forms previously ascribed to Proteacidites or Beaupreaidites from the uppermost Cretaceous and lowermost Tertiary of parts of the northern hemisphere appear to be excluded by their morphology from these genera. The fact that among the remaining forms are some that cannot at present be excluded should not lead us to the firm belief that they are Proteaceae, but only to the realisation that a degree of imprecision exists in the definition of palynomorph taxa so that they do not necessarily correspond strictly to natural families or other categories of plant taxa. Several existing families contain genera which produce Proteaceae-like pollen and in the past others may have done so. This is at least as likely as that proteaceous genera once had extensive ranges in Siberia and North America. Realisation is growing that the Late Cretaceous floras contained many large groups of angiosperms now extinct or ancestral to later forms—the unknown parent plants of the 'Normapolles', Aquilapollenites and other forms. Even in the southern hemisphere assumptions about proteaceous affinity should not go unchallenged. Total scepticism, on the other hand, may be equally unjustified. Each individual species must be considered entirely on its own, and putative affinities decided only after exhaustive comparison. Hitherto this has seldom been attempted, leading sometimes to fanciful beliefs.

REAPPRAISAL OF SOME PALYNOMORPHS 77 ACKNOWLEDGMENTS collaboration with Dr Lucy M. Cranwell, Geochronology Laboratories, University of Thanks are due to the office of Antarctic Programs, National Science Foundation, Arizona, Tucson. I am indebted to the Director of the New Washington, D.C., U.S.A., under whose grant (NSF-G-23159) work on the morphology of Zealand Geological Survey for the loan of the living and fossil Proteaceae was started in holotype of Proteacidites hakeoides. REFERENCES

COOKSON, I. C., 1950: Pollen grains of proteaceous

type from Tertiary deposits in Australia. Aust. J. Sci. Res., 3 B (2), pp. 166-177. COUPER, R. A., 1953: Upper Mesozoic and Cainozoic spores and pollen grains from New Zealand. Palaeont. Bull., Wellington, 22, pp. 1-77. COUPER, R. A., 1960: New Zealand Mesozoic and Cainozoic plant microfossils. Palaeont. Bull., Wellington, 32, pp. 1-87. ERDTMAN, G., 1966: Pollen Morphology and Plant Taxonomy. Almquist & Wiksell, Stockholm. POTONIE, R., I960: Synopsis der Gattungen der Sporae dispersae. Ill Teil. Beih. Geol. Jb., 39, pp. 1-189. SAMOILOVICH, S. R., 1967: Tentative botanicogeographical subdivision of northern Asia in Late Cretaceous time. Rev. Palaeobot. Palynol., 2, pp. 127-139. A. R. H. Martin, Botany Department, The University of Sydney, Sydney, New South Wales 2006.

Spec.Publs geoLSoc.Aust., 4: pp. 73-78, PL 1, 1973.

R., et al., 1961: Pollen and spores of western Siberia, Jurassic to Palaeocene. Trudy vses. nauchno-issled. geol.-razv. Inst., 177, pp. 1-677 (Leningrad, in Russian).

SAMOILOVICH, S.

SRIVASTAVA, S. K., 1969: Upper Cretaceous pro-

teaceous pollen from the Edmonton Formation, Alberta (Canada) and their palaeoecological significance. Can. J. Bot., 47, pp. 1571-1578.

THOMSON, P. W . , & PFLUG, H. D., 1953:

Pollen und Sporen des mitteleuropaischen Tertiars. Palaeontographica, 95 B, ( 1 - 4 ) , pp. 1-138. Two new fossil pollen genera from upper Campanian (Cretaceous) rocks of Montana. Prof. Pap. U.S. geol. Surv.,

TSCHUDY, B. D . , 1 9 7 1 :

750 B, p p . B 5 3 - B 6 1 .

A. R. H. MARTIN

EXPLANATION OF PLATE PLATE 1

All photographs of extant pollen are from slides in the University of Sydney Botany Department permanent collection. (all magnifications X 1000) Fig. 1. Protea tenox R.Br.: pollen grain 30^ diam., oblatoid, semi-angular, triporate. Sectional view; note absence of post-atrium. Fig. 2. Macadamia ternifolia F. von Muel.: pollen grain 26M- diam., oblate, sub-angular, triporate. Sectional view: note presence of post-atrium. Fig. 3. Grevillea punicea R.Br.: pollen grain 70p, diam., oblate, semi-angular, triporate, tectate-perforate, columellate; inflated pore membrane. Surface view. Fig. 4. Carnarvonia araliaefolia F. von Muel.: pollen grain 17^ diam., suboblate, semiangular, triporate, finely reticulate. Surface view. Fig. 5. Telopea speciosissima R.Br.: pollen grain 44^ diam., suboblate, semi-angular, triporate, smooth. Sectional view: note absence of post-atrium. Fig. 6. Aulax pinifolia (L.) Berg.: pollen grain 33|n diam., triporate, reticulate & verrucate. Near-sectional view; note slight ectannulus. Fig. 7. Adenanthos barbigera Lindl.: pollen grain 40m- diam., oblatoid-peroblatoid, lobatetriquetrous, triporate; reticulate, simplicolumellate. Surface view. Fig. 8. Franklandia fucifolia R.Br.: pollen grain 78n diam., spheroidal, triorate; baculate. Equatorial view. Fig. 9. Pometia pinnata Forst. (Sapindaceae): pollen grain 65|n diam., oblate, semi-angular, tricolporate; reticulate. Surface view; endannulus and vestibule show indistinctly. Fig. 10. Proteacidites sp. indescr.: pollen grain 32|LI diam., oblate, semi-angular, lobate, triporate, reticulate, pluricolumellate. Surface view (Early Tertiary coal, Comaum bore; Prep. I. C. Cookson). Fig. 11. Beauprea elegans Br. & Gr.: pollen grain, detail of aperture structure; note absence of vestibule. The short colpoid is entirely encompassed by the post-atrium-like zone —a thinning (=solution) in the inner endexine.

A.

R.

H.

MARTIN

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE

1

THE ECOLOGICAL SIGNIFICANCE OF TROPICAL MANGROVES IN THE EARLY TERTIARY FLORAS OF SOUTHERN AUSTRALIA By D. M. CHURCHILL (With 1 Table, 1 Text-Figure and 1 Plate) ABSTRACT

Tropical mangrove plant communities have been recognized as an element of the Early Tertiary pollen and wood floras from sediments of the Plantagenet Group and the Kings Park Shale. From the present-day ecology of these tropical mangroves, and their Tertiary occurrence in Australia, it is reasoned that the hottest of the coastal, wet tropics extended over a belt that reached not less than latitude 35°S and probably latitude 60°S during the Middle to Late Eocene. The tropical conditions were accompanied by a rise of sea-level, 300 m higher than at present. This produced a coastline around Western Australia that was characterized by a large number of islands, wide estuaries and extensive shallow seas. These tropical insular conditions were ideally suited for isolation and speciation of plants and animals, before they were joined once more to the mainland as the sea later receded. The height of sea-level during the Tertiary is therefore seen to have had a considerable influence on the evolution of the present Australian flora.

INTRODUCTION Pollen analysis has developed into a powerful analytical tool in palaeoecological investigations that has been applied with success to problems in Pleistocene geology, botany, zoology, archaeology and climatology. Whatever the problem however, it is always vegetation that produces the pollen which is analysed, and knowledge of vegetation is an essential requirement for the intelligent use of pollenanalytic data (Faegri, 1966). The identification of vegetation types requires some knowledge of the ecology and sociology of the vegetation concerned, and any study of past vegetation must be in error to the extent to which the ecology of the species changes. With annual plants this error could be considerable. With forest trees, where there is one generation every 250-500 years, the error should be relatively low. If the amount of change per generation is comparable, then 2,000 generations (years) of annual plant evolution is equivalent to 1 million years of forest history. Since 1925, (Turesson, 1925, 1930) ecotypes have been demonstrated to be not only genetically and physiologically stable, but closely in harmony with the habitats in which they grow (Clausen et al, 1945), and van Steenis (1962) has argued convincingly (p. 312) that; 'the world's Spec.Publs geoI.Soc.Aust., 4: pp. 79-86, PL 1, 1973.

flora is predominantly a conservative relict flora with a long history behind it, it was of very slow growth and disinclined to change', and (p. 290); 'I conclude that there is no reason to assume significant changes during the geological past in the thermo-ecology of the majority of the tropical genera concerned'. The interpretation of the Pleistocene interglacial vegetation at higher latitudes is based on plant responses to environmental changes rather than on change in the ecology of the species. These interpretations, however, carry weight because communities rather than individual species are considered; particularly when the communities in question can be correlated with well-known habitats such as mangrove or salt marsh. Because certain vegetation characterizes each of these habitats, it is necessary to define the environmental conditions as well as the vegetation. One of the greatest obstacles to the identification of Tertiary vegetation types is the paucity of pollen data from modern offshore environments at different latitudes. RECOGNITION OF VEGETATION TYPES IN OFFSHORE SEDIMENTS To carry pollen analytic vegetation studies back into the Tertiary it is important to have some idea of what a pollen spectrum from a

D. M. CHURCHILL

sample might represent in time and in space, in relation to the thickness of the sample. In uncompressed Holocene peat sediments, the pollen spectrum from a sample one cm deep contains a representative amount of pollen liberated during the diurnal, seasonal and life-span periods of many short-lived species. Within a metre, a representative amount of pollen produced by one or more communities of plant species is often preserved. Within ten metres of peat, the pollen produced by the succession of many communities can be expected. Flattening of spherical pollen is a guide to the minimal amount of compression of organic sediments. In Rollo's Bore near Coolgardie, bands of lignite one cm thick contain pollen flattened by more than 95% and therefore accumulated during the residence time of an entire community, which could be as little as 250-500 years. In space, the pollen spectrum is the integral of the pollen source vegetation, its area, distance from the sampling

site, and aerodynamic properties of the sporomorphs. The distance off-shore that pollen is found in marine inorganic sediment, in quantities sufficient for detection by routine pollen analytic procedures, was first demonstrated by Muller (1959), who found more than 10,000 grains per gram in low-latitude sediments less than 16 km from shore. Eighty km from shore the amount of pollen fell to less than 500 grains per gram. Groot & Groot (1964) and Groot et al. (1967) found between 10-100 pollen grains per gram in plain lutite sediment in 12 metres of core taken up to 2,200 km off-shore in the abyssal depths (5,000-6,000m) of the Argentine Basin. This quantity of pollen was sufficient to construct pollen diagrams that could be related meaningfully to interglacial (humid Podocarpus-Nothofagus forest and moorland) alternating with glacial (chenopod, Ephedra) vegetation. Although they recognized that the evidence

TABLE I

Numbers ofpollen grains encountered in a moss on Signy Island, South Orkneys, lat. 60° 40' S. Depths from surface (inches)* SPOROMORPH Arboreal Nothofagus N. betuloides Blume N. antarctica Oerst. N. dombeyi Blume N. pumilio Reiche N. sp. ? fusca Type Podocarpus SHRUBS Chenopodiaceae Ephedra Araliaceae Epacridaceae HERBS Plantago Compositae Umbelliferae Schizolaena Acaena Gunnera SEDGES & GRASSES Typha angustifolia Restionaceae Grass PTERIDOPHYTES Lycopodium Pteridium ALGAE Pediastrum Total * 1 inch = 2-54 cm.

0-2 26 8 1 2 20 5 22 4 14 3 2 6 4 1 1 1 1 1 37 1 1 161

6 22

1 25 8 1 8

12 18 24 30 36 48 18 6 20 19 41 10 Further identification of Nothofagus species not attempted

54 14

60 5

2 2 2

1 4

31 1 4 1

1 5 3 1

17 4 1 3

2 13 2 5

2 11 3

1 6 2

12 8

3 2

2

4

3

33

14

11

18

17

12

12

13 1

4

98

72

28

62

80

1 77

1 32

1 37

1 14

66 (Q-801 107 AD) 21

26

ECOLOGICAL SIGNIFICANCE OF TROPICAL MANGROVES was scant, Groot et al. (1967) argued that EOCENE MANGROVES the pollen in the lutites was transported with Muller (1959) demonstrated that it is posthis very fine inorganic fraction from South sible to identify tropical estuarine, riparian and America rather than Antarctica. mangrove vegetation from the pollen spectra On the Scotia arc, south of the Argentine of near-shore sediments less than 80 km from Basin, there is a 1.8 m bank of moss growing the coast. He went on to demonstrate (1964) on snow-covered Signy Island in the South an unbroken succession of tropical mangrove Orkneys, lat. 60°40'S (comparable to the lati- vegetation from the Lower Tertiary to the tude of southern Australia during the early Ter- Recent, in northwest Borneo. tiary). The base of this moss has been radioThe pollen of the monotypic genus Nypa, carbon dated (Q-801) to 107 A.D. and perma- has been described by Erdtman (1969) and frost extends to the base from 15 cm beneath by Muller (1964), and there is little chance the surface. This moss indicates the amount of of confusing it with pollen of other taxa. Nypapollen carried over long distances by wind, typo pollen (Plate 1, Figs la-c) occurs in the and the vegetation communities that can be Kings Park Shale of the Perth Basin (Clareexpected in sediments a long way off-shore. mont Asylum Bore, 183-285 ft).* Prior to Samples were collected in 1962 for pollen Muller's illustration of the pollen of Nypa, analysis and some preliminary results are given some pollen grains from the Kings Park Shale in Table I. The amount of pollen ranged had been recorded in the form taxa by through the order of magnitude 100 to 1,000 Churchill (1961) as Monosulcites perspinosus pollen grains per gram dry weight of organic Couper, of unknown affinity and therefore of matter. no ecological significance. The recognition of Although small patches of a grass—Des- these grains as Nypa immediately raises the champsia antarctica Desv., and 'pink',—Colo- possibility of other mangrove pollen types that banthus crassifolius Hook, f., (Caryophylla- have previously been ignored, or relegated to ceae), occur in small radiation traps close to ecologically meaningless form-genera. Muller (1964) doubted the identification sea-level on the northern faces of parts of the island, most of the pollen in the moss is of Santalumidites cainozoicus Cookson & Pike carried down-wind for at least 1300 km (dis- (1954). He pointed out that the Australian tance to Statten Island). That this pollen can Tertiary grains described by Cookson & Pike settle to the bottom of the ocean and become were similar to Sonneratia, a tropical mangrove incorporated in the sediments accumulating genus. Cookson & Pike were aware of the there seems as probable as its transport while difficulties of matching the Tertiary pollen attached to the fine inorganic lutite fraction with Santalum, but even now, the only desof an abyssal sediment. Clearly the presence criptions of Sonneratia pollen are those of of such pollen is not suitable evidence for Erdtman (1952), Guinet (1962) and Muller tracing the source of the lutites in the Argen- (1964), and it requires further studies of the pollen of the Sonneratia species before detine Basin. tailed comparisons with the varied Australian The work on the inorganic sediments of Tertiary species will be possible. Sonneratiathe Argentine Basin and the organic terrestrial type pollen is widespread in both the Plantadeposits of Signy Island, demonstrates that genet Group (Nornalup lignite, Munrilup Bore, pollen travels in air-streams for at least 1600 65 ft) and the Perth Basin (Rottnest Island km off-shore and can be anticipated in inor- Bore, 1285-2185 ft) (Plate 1, Figs 2-5), ganic sediments in numbers sufficient to extract and is characterized morphologically by pores, and count during routine pollen analysis. with or without an annulus and with three Furthermore it is clear from the Signy Island narrow endexinous colpi. pollen that whereas most terrestrial pollen sinks Fossil pollen grains that closely match the are dominated by nearby littoral plants such as pollen of Avicennia have been found in the Cyperaceae and Restionaceae as well as Nothosediments of the Plantagenet Group (354 mile fagus and Podocarpus (Auer, 1958), the Hay-Denmark Railway Bore, 50 ft); Plate 1, littoral species component fades into obscurity Figures 7, 8. Just how closely they also rewhen the pollen sink is removed well away semble other genera of the Verbenaceae howfrom the littoral area. ever, must await a closer study of the pollen * Imperial units have been used in bore logs, and are retained for presentation of borehole information m tnis paper. One ft = .305 m. Spec.Publs geoI.Soc.Aust., 4: pp. 79-86, PL 1, 1973.

82

D. M. CK

of that family. Fossil pollen grains that closely resemble genera in the Rhizophoraceae, Rhizophora or Ceriops, have also been found in the Plantagenet Group (Munrilup Bore, 65 ft) and from Coolgardie (Rollo's Bore, 390 f t ) ; Plate 1, Figure 6. These are tricolporate grains with a very elongated endexinous furrow at right angles to the colpi. It is clear that a more detailed study of pollen of Malesian families is necessary before substantial advances can be made in the interpretation of the Australian Tertiary and Late Cretaceous angiosperm strandline floras. However from the evidence presented above, tropical mangrove vegetation was certainly a component of the Middle to Late Eocene floras of southwestern Australia. SUPPLEMENTARY EVIDENCE The ecology of extant mangrove vegetation has been summarized by Hou (1958), Richards (1957), Ridley (1930) and van Steenis (1962). Mangrove vegetation occurs along coastlines that fringe shallow seas where there are large tidal rivers that transport a regular supply of silt from inland granitic ranges. Mangroves do not occur along arid shores, due to the absence of rich silt-carrying rivers, and they do not occur at the outfall of rivers where the inland geology is sandstone and the littoral shores are sandy. Tidal mud implies a continent or large island with large rivers to bring down the class of silt which forms the characteristic blue or grey mud in which mangroves grow. The development of mangrove vegetation follows the silting up of coastal areas but does not initiate the process. The lithology of the Kings Park Shale (calcareous organic shales) and the Toolbrenup beds (silicified mudstones and sandstones) in the Plantagenet Group is certainly compatible with the type of environment outlined above. It is not difficult to envisage the occurrence of Nypa—Rhizophoraceae—Sonneratia—A vicennia mangrove vegetation on the muddier of these facies. The presence of abundant, well-preserved silicified Barringtonia wood (Plate 1, Figs 9-13) in the Toolbrenup beds of the Plantagenet Group (identified by Mr H. D. Ingle, Forest Products Laboratory, C.S.I.R.O.) suggests that Barringtonia sandy beaches may have been interspersed with the mangroves along the strandlines of the MiddleLate Eocene shores of southwestern Australia. The temperatures of water in which these

strandline communities grow today, have minimum values not less than 20-25°C. If the thermal ecology of these strandline communities has not changed throughout the Tertiary then the minimum temperatures of the in-shore coastal waters of southwestern Australia were at least 5-10°C warmer than at present, if Australia was at its present latitude. Thus, the fossil pollen and wood evidence indicates that tropical coastal waters extended along these southwestern and southern shores during the Middle to Late Eocene, at whatever latitude Australia occupied at the time. PHYSICAL CHARACTERISTICS OF COASTAL ENVIRONMENTS OF WESTERN AUSTRALIA DURING T H E EOCENE The latitude of Australia during the Early Tertiary, determined from palaeomagnetic work by Irving (1967), appears to have been much farther south than its present position, with the southern Australian mainland at about lat. 60°S (Fig. 1). The Precambrian shield of Western Australia has had no isostatic load from above since the Permian glaciation and it therefore provides an ideal reference level for eustatic changes of sea-level. The presence of widespread horizontal, unfaulted marine sediments, of post-Permian age, at different levels on the shield may be taken as reliable evidence of the minimal height of the contemporary sea-level relative to present sea-level. The lithological evidence of marine transgression during the Eocene in Western Australia is as striking as that found during the course of the Flandrian transgression of the Holocene; namely coastal freshwater peat beds that formed close to the contemporary sea-level have been overlain by marine sediments of the transgression (Churchill, 1965; Jelgersma, 1961; Godwin, 1943). The Pleistocene levels of the sea are controlled by temperature which affects the degree to which the polar ice caps have melted and discharged the melt-water into the sea. The sediments of the Plantagenet Group include isolated freshwater lignite beds that overlie a basement of shallow sands or granite, but are overlain by extensive, unfaulted, horizontally bedded, marine shales, siltstones, claystones, sandstones and limestones. Some facies of these marine beds are richly fossiliferous. Levels at which freshwater lignites have been overtaken by Eocene marine sediments range from 46 m below sea-level (Werillup Bore-

ECOLOGICAL SIGNIFICANCE OF TROPICAL MANGROVES hole) through —15 m (Hay River Bore) + 3 m (Nornalup lignite) +43 m (Albany Bore No. 15), +60 m (Shark Lake near Esperance), +207 m (No. 6 Bore, Munrilup), +244 m (Lake Cowan, Norseman). The localities and stratigraphic details of these bores may be found in Maitland (1901) and Cockbain (19686), and with the exception of the Weril-

Spec.Publs geol.Soc.Aust, 4: pp. 79-86, PL 1, 1973.

83 lup Bore, all have been analysed for pollen (Churchill, unpublished data). It is not known whether the beds that overlie the lignites at about 290 m (Rollo's Bore) near Coolgardie are marine or freshwater as only the lignites were analysed for pollen (Balme & Churchill, 1959). At Lake Cowan, Norseman, however the marine conditions that overtook the fresh-

D. M. CI water phase, progressed to the deposition of the than they are today, then there is conflict beshallow-water inshore Norseman Limestone, tween the palaeomagnetic and palaeobotanical the Cowan Dolomite and finally the off-shore evidence. However, the occurrence of Nypa Princess Royal Spongolite, as the marine trans- and many other tropical Indo-Malayan elegression moved even farther inland. The ments in the Eocene London Clay flora, from marine beds of this sequence have been named lats 50°-52°N in England (Chandler, 1964), the Eundynie Group, and the Foraminifera suggests that the tropics did occupy a belt that from the Norseman Limestone are of Late was some two to three times wider than at present. The Late Eocene marine transgression Eocene age (Cockbain, 1968a). It is of considerable biological and palaeo- 300 m on to the southern edge of the continent geographic significance to reconstruct the dis- from Albany inland to Kojonup, and Espertribution of land above sea-level, when sea- ance inland to Norseman, provides a horilevel was 300 m higher than at present, as it zontal datum against which to measure the was (to within 60 m) during the maximum of bathymetry and approximate shoreline conthe Late Eocene marine transgression. Figure 1 figuration of the period (Fig. 1). Even if the shows the present outline of Australia and the present 300 m (1,000 ft) contour is in error, present 300 m (1,000 ft) contour. When sea- relative to that shoreline by as much as 60 m level was 300 m higher than at present, the it is evident from Figure 1, that the Western 1,000 ft contour would represent the old Australian coastline was fringed by a very shore-line. However, as a shore-line map, it is large number of islands that would have proin error in the eastern states to the extent vided physical conditions ideal for the geothat the Middle to Late Tertiary epeirogeny graphic isolation necessary for speciation of raised more land above sea-level than would insular floras and faunas. These tropical conhave been showing during the Eocene. Never- ditions would have been analagous to the theless, the insular nature of the region, the Galapagos Islands today. By contrast, when large number of offshore islands, estuaries, sea-level was only at the 150 m (500 ft) and the extensive shallow seas during Late contour, it is evident from Figure 1, that few if any of these islands would have existed. Eocene times are at once apparent. Allowing only half the two million year interval generally credited to have elapsed through the Middle and Late Eocene, for isoDISCUSSION lation of the flora on offshore islands during The Late Eocene occurrence of Nypa, the the marine transgression, this is equivalent to tropical mangrove palm, Sonneratia, Avi- about 20,000 generations of forest trees. Whether the large number and concentracennia and the Rhizophoraceae mangroves, and Barringtonia along the southern margins tion of species of Casuarina, Banksia, Dryanof Western Australia, carries many implica- dra, Hakea and Grevillea found in southtions. That this type of vegetation did not western Australia today, achieved reproductive originate in a temperate climate and invade isolation under the tropical insular conditions the tropics during the Tertiary is proved by prevailing at that time, may never be proved. the occurrence of Nypa, the Rhizophoraceae Certainly they are well represented in the fossil and Sonneratiaceae mangroves throughout the pollen and wood floras of the Eocene, alongperiod in Borneo, whereas there has been a side the mangrove elements described above, loss of these elements from southern Australia and they have survived to the present day. Other elements of these tropical floras, ranging since the Eocene. The geographical position of southern Aus- from Nothofagus to Anacolosa-type Olacaceae tralia during the Late Eocene was as far south and the mangroves themselves, with the excepas lat. 6 0 ° S , on palaeomagnetic evidence. If tion of Avicennia, became extinct; presumably the equator is assumed to have occupied its due to the shift in the tropics by contraction present position, and the tropics were no wider or angular rotation. 8 4

REFERENCES

AUER, V., 1958: The Pleistocene of Fuego-Pata-

gonia. Part II: The history of the flora and vegetation. Suomal. Tiedeakat. Toim. A. 111., 50.

BALME, B. E., & CHURCHILL, D. M., 1959: Ter-

tiary sediments at Coolgardie, Western Australia. /. Proc. R. Soc. West. Aust., 42, pp. 37-43.

ECOLOGICAL SIGNIFICANCE OF TROPICAL MANGROVES 85 CHANDLER, M. E. J., 1964: The Lower Tertiary GROOT, J. J., GROOT, C . R., EWING, M . , BURCKLE, Floras of Southern England, 4. A Summary L., & CONOLLY, J. R., 1967: Spores, pollen,

of Findings in the Light of Recent Botanical diatoms and provenance of the Argentine Observations. Brit. Mus. Nat. Hist., London. Basin sediments. Prog. Oceanog., 4, pp. 179-217. CHURCHILL, D. M., 1961: The Tertiary and Quaternary vegetation and climate in relation P., 1962: Pollens D'Asie Tropicale. to the living flora in South Western Aus- GUINET, Inst. fr. Pondichery Trav. Sec. Sci. Tech., tralia. Ph.D. Thesis, Univ.West.Aust. (unpub(1). lished). CHURCHILL, D. M., 1965: The displacement of Hou, Ding, 1958: Rhizophoraceae. Flora Malesiana., 1, pp. 429-493. deposits formed at sea-level, 6,500 years ago in southern Britain. Quaternaria, 7, pp. IRVING, E., 1967: Palaeomagnetic evidence for 239-249. shear along the Tethys; in Aspects of Tethyan CLAUSEN, J., KECK, D . D . , & HIESEY, W . M . , Biogeography. Pubis Syst. Ass., 7, pp. 59-76. 1945: Experimental studies on the nature of JELGERSMA, S., 1961: Holocene Sea-level Changes species. Pubis Carnegie Inst., 564. in the Netherlands. Vitjeversmij E. van COCKBAIN, A. E., 1968a: Eocene Foraminifera Aelst., Maastricht. from the Norseman Limestone of Lake Cowan, Western Australia. Rep. geol. Surv. MAITLAND, A . G . , 1901: Boring for coal near West. Aust. for 1967, pp. 97-98. Albany. Rep. geol. Surv. West. Aust. for 1900, p p . 16-20. COCKBAIN, A. E., 19686: The stratigraphy of the Plantagenet Group, Western Australia. Rep. MULLER, J., 1959: Palynology of Recent Orinoco geol. Surv. West. Aust. for 1967, pp. 99-101. delta and shelf sediments. Micropaleontology, COOKSON, I. C., & PIKE, K . M . , 1954: Some 5, pp. 1-32. dicotyledonous pollen types from Cainozoic deposits in the Australian region. Aust. J. MULLER, J., 1964: A palynological contribution to the history of the mangrove vegetation in Bot., 2, pp. 197-219. Borneo; in Cranwell, L. M. (Ed.), Ancient ERDTMAN, G., 1952: Pollen Morphology and Pacific Floras, pp. 33-42. Univ. Hawaii Press, Plant Taxonomy. Almqvist & Wiksell, StockHonolulu. holm. ERDTMAN, G . , 1969: Handbook of Palynology. RICHARDS, P. W., 1957: The Tropical Rain Forest. Cambridge Univ. Press, London. Hafner, New York. FAEGRI, K., 1966: Some problems of representivity RIDLEY, H . N . , 1930: The Dispersal of Plants throughout the World. Reeve, London. in pollen analysis. Palaeobotanist, 15, pp. 135-140. TURESSON, G., 1925: The plant species in relation GODWIN, H., 1943: Coastal peat beds of the to habitat and climate. Hereditas, 6, pp. British Isles and North Sea. /. EcoL, 31, pp. 147-236. 199-247. G., 1930: The selective effect of climate GROOT, J. J., & GROOT, C. R., 1964: Quaternary TURESSON, upon plant species. Hereditas, 14, pp. 99-152. stratigraphy of sediments of the Argentine Basin: a palynological investigation. Trans. van STEENIS, C . G. G. J., 1962: The land-bridge N.Y. Acad. Sci., 26, pp. 881-886. theory in botany. Blumea, 11, pp. 235-372. D. M. Churchill, Department of Botany, Monash University, Clayton, Victoria 3168.

Spec.Publs geol.Soc.Aust., 4: pp. 79-86, PI. 1, 1973.

Present address: Royal Botanic Gardens, South Yarra, Victoria 3141.

D. M. CHURCHILL

EXPLANATION OF PLATE

PLATE

Fig. 1. (a-c). Nypa-type

1*

pollen. Claremont Asylum Bore, 183-285 ft.

Figs. 2, 3, 5. Sonneratia-type pollen. Three grains from the Albany Bore No. 6, Munrillup, 65 ft. Fig. 4. Sonneratia-type pollen. Rottnest Island Bore, 1,595-2,021 ft. Fig. 6. Rhizophoraceae-type pollen. Albany Bore No. 6, Munrillup, 65 ft. Figs. 7, 8. Avicennia-type pollen. Hay-Denmark Sample 8662/1. 50 ft.

Railway Bore, 354 miles from Perth.

Fig. 9. Barringtonia. Silicified wood. Toolbrenup beds. Spec. 2503T. L.R.S. Vessel adjacent to (arrow) medullary ray. Note elongation of pits adjacent to ray cells, and the polygonal form (above arrow) of the pit outlines, away from the ray. Fig. 10. Barringtonia. Silicified wood. Toolbrenup beds. Spec. 2503T. L.T.S. Pit field on vessel wall. Fig. 11. (a). Barringtonia calyptrata. Wood from trees living in N. Qld. C.S.I.R.O. Reference Spec. No. 8436. L.R.S. Vessel adjacent to medullary ray with elongated pit field. Away from the ray, the pits are polygonal in outline (arrow), (b). An enlargement of the polygonal pit field from the position of arrow in 11 (a). Fig. 12. Barringtonia cf. calyptrata. Wood from trees living in Reference Spec. No. 11,590. Transverse section of wood. Fig. 13. Barringtonia. Silicified wood. similar to Fig. 12.

Toolbrenup

beds. Transverse

N.

Qld.

section

C.S.I.R.O. of

wood,

Scales: 1-8, x 700, 9-10, x 300. 11(a), 12, 13, x 700.

•All slides of fossil material are housed in the National Herbarium, Melbourne. Reference slides from living wood are in the collection of C.S.I.R.O. Forests Products Laboratory, Melbourne.

D.

M.

CHURCHILL

Spec.Publs geoI.Soc.Aust., 4, 1973.

PLATE

1

SPORE ZONATION AND SEDIMENTARY HISTORY OF THE NEOCOMIAN, GREAT ARTESIAN BASIN, QUEENSLAND By D. BURGER (With 5 Tables, 10 Text-Figures and 2 Plates) ABSTRACT

The basal part of the Lower Cretaceous palynological sequence in the Great Artesian Basin, Queensland, is characterized by an overlap in the ranges of Murospora florida and Cicatricosisporites australiensis and is mainly Neocomian age. It has been widely recognized in Australia and is here designated as the Murospora florida Zone. Three local subdivisions of the zone are distinguished; they are designated in stratigraphic order as the Cicatricosisporites australiensis Subzone (?Tithonian to ?Berriasian), the Foraminisporis wonthaggiensis Subzone (middle Neocomian, Valanginian-Hauterivian?) and the Foraminisporis asymmetricus Subzone (upper Neocomian (Barremian) to basal Aptian). The limits of each of the subzones are defined by the ranges of selected spore species. The sedimentary history of the basin is characterized by an initial period with accumulation of nonmarine, porous (often aquiferous) sandstones (Mooga Sandstone, Lower Hooray Sandstone), followed by a period with generally quieter lacustrine and fluviatile deposition (Upper Hooray Sandstone, Bungil Formation), while contemporaneous marine argillaceous developments (Doncaster Member) towards the north marked an initial invasion of the sea. A third period with alternating marine and freshwater sedimentation (Bungil Formation, higher Upper Hooray Sandstone) was ended when the sea flooded the entire basin in Queensland and adjoining regions during the late Aptian. INTRODUCTION The present stratigraphic and palaeontological knowledge of the Lower Cretaceous has its roots in Europe, where detailed studies of richly fossiliferous sediments in Switzerland and France during 1840-1870 by d'Orbigny, d'Archiac, Leymerie, Coquand and others led to a subdivision of the Lower Cretaceous into nowadays internationally accepted geological stages. The Valanginian in Switzerland, and the Berriasian in southeastern France were accepted as the lowermost Cretaceous. Study of the ammonite succession within the Berriasian at its type locality (near Berrias) led Kilian and Mazenot (1939) to suggest that the Jurassic-Cretaceous boundary be placed between the upper Tithonian, with Berriasella chaperi, B. ayziensis, Dalmasiceras djanelidzei and Neocomites suprajurensis, and the lower Berriasian, with Berriasella paramacilenta and B. grandis. This boundary was also proposed at the 'Colloque sur le Cretace inferieur en France 1963', although with slight reservations, as the basal strata of the Berriasian at its type locality are poor in ammonites (Busnardo, le Hegarat & Magne, 1965). In Australia, study of the Lower Cretaceous in comparable detail has not been possible

because of the relative lack of fossils. Whitehouse (1955), for instance, in his account on the geology of the Great Artesian Basin in Queensland, noted that palaeontological evidence below the marine Cretaceous (Aptian) was virtually non-existent. Australian palaeontologists usually apply the term Neocomian, a collective term which includes the Berriasian, Valanginian, Hauterivian and Barremian (Brinkmann, 1959). In the Great Artesian Basin, where deeply weathered sediments in outcrop have been a negligible source of pollen and spores, the acquisition of sub-surface material from commercial and scientific drilling has opened a wide field for palynology. Age estimations of Jurassic and Lower Cretaceous palynological divisions proposed by Dettmann (1963), de Jersey & Paten (1964), and Evans (1966a) partly rested on Balme's (1957) deductions for the age of equivalent intervals (his Microfloras I, II-A, II-B) in the Perth, Canning and Carnarvon Basins. Evans (1966a-c) recovered his palynological unit Kla mainly from sediments below the marine Aptian and assumed, on the basis of Balme's work, that the base of the unit coincided approximately with the Jurassic-Cretaceous boundary. Dettmann (1963) and Dettmann & Playford

Spec.Publs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

D. BURGER

142°

Fig. 1.

148°

Location of drilled sections in the Great Artesian Basin.

154*

SPORE ZONATION AND HISTORY OF NEOCOMIAN

(1969) thought that their Crybelosporites stylosus Zone ranged in age from possibly uppermost Jurassic to Valanginian. The results of further detailed study of palynological unit Kla, partly on the basis of new material, are reviewed in the present paper. Also, palynological aspects of the Jurassic-Cretaceous boundary are discussed. The unit is here formally designated as the Murospora fiorida Zone. As was noted above, palaeontological evidence for the age of the associated sediments is scarce and fragmentary. A brief summary of these sediments is given below, together with the evidence for their age. NOMENCLATURE AND AGE OF ROCK FORMATIONS Late in the nineteenth and early in the twentieth century field mapping was initiated by the interest in the occurrence of oil, natural gas and artesian water in various parts of the Great Artesian Basin. Whitehouse (1955) was the first to treat the geology of the Queensland portion of the basin as a whole. He introduced a uniform terminology for the Jurassic and Cretaceous rock sequence, which was widely adopted (Geological Survey of Queensland, 1960; Union Oil Corporation et al., 1964; Cundill, Meyers & Associates, 1967). Subsequent field work in the Surat Basin (Day, 1964; Exon, Milligan, Casey & Galloway, 1967; Thomas & Reiser, 1968) with the aid of air photographs and extensive subsurface information from deep wells brought forth successive refinements of Whitehouse's nomenclature. The terminology accepted in the present paper is derived from Day (1964), Exon et al. (1967), and Exon & Vine (1970). Underneath the marine (Aptian) Wallumbilla Formation two most probably Cretaceous units are distinguished, the Mooga Sandstone and the Bungil Formation, consisting of the Kingull, Nullawurt Sandstone, and Minmi Members (Fig. 10). In the Eromanga Basin (Mitchell, Tambo, Blackall and Augathella Sheet areas) Exon, Galloway, Casey & Kirkegaard (1966) and Exon et al. (1967) mapped a sandy unit (Hooray Sandstone) underneath and in conformable contact with the Rolling Downs Group (Figs 2, 3). Exon (1966) correlated the Hooray Sandstone with the Gubberamunda Sandstone to the Bungil Formation (Fig. 10). Towards the centre of the basin the Hooray Sandstone was recognized in subsurface of

89

exploration wells and water bores (Gregory, Senior & Galloway, 1967; Senior, Galloway, Ingram & Senior, 1968; Senior, Ingram, Thomas & Senior, 1969; Senior, 1969). Here, two major divisions were recognized (Gregory et al., 1967; Evans, 1966d). The Upper Hooray (the Transition Beds' of many exploration company reports) might be the correlate of Freytag's (1966) 'Cadna-Owie Formation' in South Australia. The Lower Hooray consists in many wells of a lower and upper sandy member, separated by a thin finer grained interval (Fig. 5). Eastwards, near the outcrop of the formation, a twofold division, separated by an unconformity exists in the Tambo and Augathella areas (Woolley, 1941; Exon, 1966; Exon et al., 1966). This subdivision does not correspond to that in the central part of the basin and is probably a local phenomenon (Fig. 2). Reynolds (1960), Vine & Jauncey (1962), Vine, Casey & Johnson (1964), and Vine, Jauncey, Casey & Galloway (1965) mapped different sequences farther north in the basin. The Ronlow Beds (Fig. 3) occur between the Triassic and the Aptian in the Tangorin, Muttaburra and Galilee Sheet areas (Vine et al., 1965). In the Hughenden Sheet area two units were mapped, the Blantyre Sandstone and the Gilbert River Formation, separated by a regional unconformity (Vine et al., 1964). They lie between Triassic sediments and basement metamorphics and the marine Rolling Downs Group (Fig. 6). The Gilbert River Formation extends onto the Gilberton and Richmond Sheet areas (Grimes & Smart, 1970). At the western margin of the Eromanga Basin (Boulia and Duchess Sheet areas) Casey, Reynolds, Dow, Pritchard & Paten (1960) described the Longsight Sandstone in outcrop between basement and the marine Rolling Downs Group. Towards the centre of the basin this sandstone merges in subsurface with a broader arenaceous sequence, which Jauncey (1964) and Vine (1964a, b) referred to as 'Mesozoic undifferentiated' and which also occurs in Ooroonoo No. 1 (Fig. 6).

Whitehouse (1955), Day (1964), Vine & Day (1965), Day (in Exon et al., 1967) recorded Aptian 'Roma' faunas in outcrop of the Doncaster Member (Wallumbilla Formation). Similar faunas occur in outcrop of the Longsight Sandstone (Hill, in Casey et al., 1960;

Spec.Publs geoI.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

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SPORE ZONATION AND HISTORY OF NEOCOMIAN Reynolds, in Hill & Denmead, 1960). Laing & Power (1959) dated the Gilbert River Formation as Aptian, based on marine faunas collected from the formation near Croydon, farther north. From evidence of fossil plants White (1963) suggested a Jurassic to Early Cretaceous age for the Blantyre Sandstone. Exon (1966) and Exon et al. (1967) inferred a Late Jurassic to Early Cretaceous age for the Hooray Sandstone in the eastern Eromanga Basin, based on its stratigraphic position; there is no faunal evidence for the age of the unit. In the northern Surat Basin (Roma and Mitchell Sheet areas) evidence from below the Rolling Downs Group is also insufficient, in spite of generally better outcrop. Day (1964) thought that the 'Blythesdale Flora', containing Taeniopteris spatulata, Oleandridum jaculi, Phyllopteroides feistmanteli, P. lanceolata, Equisetites cf. E. rajmahalensis, Sphenopteris sp., S. flabellifolia, could be as old as, or slightly younger than, the Upper JurassicNeocomian Stanwell Coal Measures in the Rockhampton area. From the occurrence of cf. Phyllopteris lanceolata in the top of the Mooga Sandstone, east of Roma, and Hausmannia wilkinsi and Elatocladus planus in the Nullawurt, northwest of Roma, White (1967) suggested an Early Cretaceous age for both formations. The oldest known marine Cretaceous fauna occurs in the Nullawurt Sandstone Member northeast of Mitchell. It contains freshwater pelecypods and marine forms such as Meleagrinella sp. Day (1969) suggested that this fauna could be of Neocomian age; related Meleagrinella forms occur in the Laura Basin together with a probably Neocomian ammonite, Hatchericeras lakefieldense. A probably conspecific pelecypod (Pseudomonotis sp.) also occurs in the Neocomian of the Stanwell area. Faunas f r o m the Minmi Member in the Roma and Mitchell areas were dated as Aptian from their general resemblance to the Roma faunas (Day, 1964, 1969). SAMPLING O F F O R M A T I O N S Samples from the Orallo Formation, Mooga Sandstone and Bungil Formation in deep wells (cores, sidewall cores), as well as core samples from these formations in shallow stratigraphic bores drilled more recently, initially examined by Dr P. R. Evans and myself, were critically re-examined for the present study. Samples from the Hooray Sandstone are

93

generally poor in plant microfossils. Those selected for the present study (cores, sidewall cores) are regarded as the most reliable source of information to date (Figs 2 - 5 ) . Cores and sidewall cores from the Ronlow Beds, Gilbert River Formation and their correlate strata farther west gave satisfactory results, but palynology of the Late Mesozoic in the northern Eromanga Basin is still in an early stage. Correlation of the Lower Cretaceous sediments is based on information from four bores only (Fig. 6). More accurate data will become available with progressing work in the Carpentaria Basin. Reference to stratigraphic information of various oil exploration wells is listed in Appendix A. Stratigraphy of shallow bores was treated in unpublished records issued by the Bureau of Mineral Resources, Canberra. Bores and wells mentioned in this paper are plotted on a general location map (Fig. 1). Specifications of samples examined are given in Appendix B. PREVIOUS PALYNOLOGICAL WORK The first comprehensive study of the Lower Cretaceous in the Great Artesian Basin was undertaken by Dettmann (1963), who distinguished in subsurface of the South Australian part of the basin, as well as in southern Australia, in stratigraphic order the Stylosus, Speciosus and Paradoxa Assemblages. The Stylosus Assemblage, Dettmann & Playford's (1969) Crybelosporites stylosus Zone, is characterized by the occurrence of C. stylosus and Aequitriradites hispidus. This zone, together with part of the Speciosus Assemblage, or Dettmann & Playford's (loc. cit.) Dictyotosporites speciosus Zone, which is characterized by the presence of the nominate species and the absence of C. stylosus, are associated with the nonmarine Cretaceous sequence. Subsequent studies of the Lower Cretaceous of the basin in Queensland by Evans led to a provisional subdivision of the palynological sequence into units, which were traceable throughout Queensland and Papua. The lowermost unit K l a occurred mainly underneath the marine Rolling Downs Group in Queensland, and was also recorded in the Papuan and Morehead Basins (Evans, 1966a, e; Burger, 1968, 1969; also unpublished information). The unit is characterized as that interval in which Murospora florida and Cicatricosisporites australiensis occur together, and

Spec.Publs geaLSoc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

F D.S. INNAMINCKA No I

5.

SMART OIL ORIENT No 2 2

ALLIANCE CHANDOS Nol 3

PHILLIPS-SUNRAY COTHALOW No.l 4

ALLIANCE YONGALA Nol 5

W.O.L.No3 W.O.L.No2 (WARBRECCAN) 6

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Schematic profile through the central and southern Eromanga Basin. For additional explanation of symbols see also Figure 3.

SPORE ZONATION AND HISTORY OF NEOCOMIAN CONORADA

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OOROONOO No I

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I

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Doncaster Member Gilbert River Formation Blantyre Sandstone Loth Formation

Fig. 6. Schematic profile through the northern Eromanga Basin. For additional explanation of symbols see also Figure 3. is as such comparable to Balme's (1957) Neocomian-Aptian Microflora II-B. From its stratigraphic position, the presence of C. stylosus, and the general resemblance of the microfloras it was clear that the unit was at least in part equivalent to the Crybelosporites stylosus Zone. Further comparison proved difficult; A. hispidus is virtually absent in Queensland and Papua, and C. stylosus is infrequent to rare in Queensland, and is very rare in Papua (Evans, 1966e; also unpublished information). The Crybelosporites stylosus Zone is therefore not satisfactory for stratigraphic application in those areas. The concept of unit K la has proved to be more reliable; with added study of new material from Queensland the unit could be further subdivided, on the basis of first occurrence of species. The results are discussed in detail below. THE PALYNOLOGICAL ZONATION The microfloral assemblages recovered from the nonmarine part of the rock sequence

(sandstones, siltstones) are often poorly preserved as they have been affected in various degrees by deep weathering of the sediment; identification of specimens is thus sometimes only tentative. The occurrence of palynomorphs, also those of uncertain affinity, is documented in Tables I-III*. Reworked elements, mainly from the Permian, were found in many microfloras; they are not included. Vertical ranges and bibliographic references of species are given in Tables IV and V. Microscope preparations containing the assemblages studied are kept in the Palynological Laboratory of the Bureau of Mineral Resources. Preparations which contain the specimens illustrated in Plates 1 and 2 are stored in the Commonwealth Palaeontological Collection in Canberra. The illustrated specimens are provided with a Collection Number (CPC): their locations in the preparations are expressed in vernier readings for Leitz Ortholux Binocular microscope No. 741826, used in the Palynological Laboratory.

* All tables are set out at the end of this paper. Spec.Publs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

96

D. Bl Cyathidites australis, C. minor, Gleicheniidites fiorida ZONE Lycopodiumsporites austroclavaCharacteristics: The Murospora fiorida Zone, circinidites, Osmundacidites wellmanii, Stereias well as its subdivisions, are defined by the tidites, antiquasporites, Vitreisporites pallidus. occurrence of selected index species. They sporites Regularly but much less abundantly occurring are not tied to any rock type. Each interval is species are: Alisporites grandis, Araucariacites indicated by the name of one of the index australis, Callialasporites dampieri, Ceratospecies; however, it should be understood that equalis, Classopollis spp., Contignithis species is not necessarily present in every sporites sporites cooksonii, mortoni, microfloral assemblage within that interval, Foraminisporis dailyi,Dictyophyllidites Leptolepidites verrucaor even restricted to it (Teichert, 1958). tus, Lycopodiumsporites rosewoodensis, I consider these intervals to be equivalent or antarcticus, Neoraistrickia approximately equivalent in time, wherever Microcachryidites Polycingulatisporites densatus. they occur, in concurrence with the principle truncata, Species which are or less common in by which Dettmann & Playford (1969) charac- stratigraphically lowermoremicrofloras also terized their Cretaceous zones. In the marine persist into the zone and succeeding but Rolling Downs Group there are more direct although sometimes sporadically, are:intervals, Annucriteria to indicate isochrony of pollen boun- lispora folliculosa, Callialasporites trilobatus, daries, such as parallel relationship with dis- Cingutriletes clavus, Inaperturopollenites turtinct faunal horizons. Furthermore, the resem- batus, Staplinisporites caminus, Trilobosporites blance between palynological sequences des- antiquus, Tsugaepollenites segmentatus. cribed from the Lower Cretaceous in the Species occurring sparsely zone and Otway, Gippsland and Great Artesian Basins adjacent intervals, frequently inas the a single specistrongly suggests uniform evolution of the men are: Aequitriradites spinulosus, A. verruvegetation throughout eastern Australia. cosus, Dictyotosporites complex, lschyoThe Murospora fiorida Zone is the equiva- sporites punctatus, Leptolepidites major, Lycolent interval of Evans' (1966a) palynological podiumsporites eminulus, L. reticulumsporites, unit K la. It succeeds his (Jurassic) units J Monosulcites minimus, Podocarpidites multe5-6 and is in turn succeeded by his (Aptian) unit K lb-c. The zone is defined as the inter- simus. The following species have been found val which commences with the first appearance within zone and in younger intervals, but of Cicatricosisporites australiensis, C. hughesi, are notthedefinitely identified from the precedCouperisporites tabulatus, and Cyclosporites ing unit J 5-6: Biretisporites potoniaei, Cyahughesi, and which contains Murospora thidites concavus, Reticulatisporites pudens, fiorida and Nevesisporites vallatus. M. fiorida Trilites cf. T. tuberculiformis, and Velosporites also occurs in comparable frequency in the triquetrus. So far, evidence indicates that the Upper Jurassic of the Great Artesian Basin first occurrence of Reticulatisporites pudens (Evans, 1966a). It has not been observed in and Trilites cf. T. tuberculiformis may apyounger than Aptian strata; the evidence sug- proximately coincide with that of Cicatricosigests that it disappears from the sequence sporites australiensis. The zone contains prior to the development of the late Aptian furthermore the initial appearances strati'Roma' faunas. N. vallatus occurs in the graphically important species such as ofCrybeloJurassic of the Surat and Eromanga Basins sporites stylosus, Rouseisporites simplex, and its stratigraphically highest occurrence linearis, K. majus, Dictyoappears to coincide approximately with that of Kraeuselisporites tosporites speciosus, Foraminisporis wonthagM. fiorida. Crybelosporites punctatus Dett- giensis, mann, 1963, and Osmundacidites cf. O. mollis notensis. F. asymmetricus, and Pilosisporites (Cookson & Dettmann), which occur in unit K lb-c and succeeding intervals, have not Occurrence: Standard for the zone is taken as been observed in the zone. Further detailed the interval between core 40 at 406 feet (123.5 study, based on new material, is under way to m) and core 16 at 185 feet (56.4 m) in BMR determine the upper limit of the zone more Roma No. 1 Scout Hole, Surat Basin. This inaccurately in the Great Artesian Basin. terval includes the Mooga Sandstone and the Common and frequently occurring species Bungil Formation (Kingull Member, Nullawurt Sandstone Member, and basal Minmi within the zone as well as adjacent parts of the sequence are: Alisporites similis, Baculati- Member). Further occurrences of the zone are given in Tables I to III. sporites comaumensis, Concavisporites spp.,

THE Murospora

SPORE ZONATION AND HISTORY OF NEOCOMIAN 97 Age: Of the species that first appear at the of which 49 represent the largely unchanged lower limit of the zone, most attention in recent 'Jurassic' element. This constitutes a net inyears has been given to Cicatricosisporites aus- crease of more than 30 species, which is more traliensis and closely related forms, because of than 50% of the number of species in the their global occurrence in the Lower Cre- Upper Jurassic. This increase includes many taceous and apparent absence from all but species which Cookson & Dettmann (1958) the uppermost Jurassic. C. australiensis and and Dettmann (1963) described from the morphologically related forms (C. mohrioides, lowermost Cretaceous in the Great Artesian C. dorogensis, C. abacus) have indeed been Basin (South Australia), the Otway and found in uppermost Jurassic sediments in Gippsland Basins, and which they recognized Canada (Pocock, 1962), Britain (Lantz, as being stratigraphically significant. 1958; Norris, 1969) and Holland (Burger, On the basis of restricted ranges of these 1966). species, three sub-units are distinguished, In the Canning Basin, Western Australia, which are here designated, in stratigraphic Balme (1957) encountered forms related to order, as the Cicatricosisporites australiensis C. australiensis in Broome No. 3 Bore, in strata Subzone, the Foraminisporis wonthaggiensis which Veevers & Wells (1961) on faunal Subzone, and the Foraminisporis asymmetrievidence regard as Upper Jurassic, probably cus Subzone. I do not rank these units as Oxfordian or Kimmeridge. He also observed zones, as it is not known whether they are C. australiensis in the Jarlemai Siltstone, dated identifiable in the Lower Cretaceous outside as Oxfordian to ?Tithonian in the same basin, the Great Artesian Basin. as well as related forms in Tithonian beds in subsurface of Barrow Island, Carnarvon THE Cicatricosisporites australiensis SUBZONE Basin (unpublished information cited by Dett- Characteristics: The Cicatricosisporites ausmann & Playford, 1969, pp. 186-7). In eastern traliensis Subzone is the lowermost of three Australia it is at present not possible to assess subzones which together form the Murospora the earliest occurrence of C. australiensis by florida Zone. It is distinguished from the premeans of direct palaeontological evidence. ceding sequence by the presence of C. ausIn the Papuan Basin, Evans (1966e) re- traliensis, C. hughesi, Couperisporites tabucorded a unit K la microflora from APC latus, and Cyclosporites hughesi. The followlehi No. 1 at 4,712 feet (1,436.2 m), while ing species, Biretisporites cf. B. potoniaei belemnites, dated as late Neocomian by Cyathidites concavus, Reticulatisporites Glaessner (in Gay & Brown, 1961), occurred pudens, Trilites cf. T. tuberculiformis, and at 4,717-9 feet (1,438 m) and at 4,745-6 feet Velosporites triquetrus, which occur more or (1,466 m). In the Surat Basin the zone occurs less infrequently throughout the subzone, have in the Minmi Member, which, as we have not been definitely identified in older microseen, contains early Aptian faunas in out- floral assemblages, although their presence crop. Evans (1966a) recorded the succeed- prior to the subzone is at present not coning unit K lb-c from the late Aptian Don- sidered impossible. Furthermore, the subzone caster Member (Wallumbilla Formation). includes the earliest occurrence of AequitriraOn the basis of this evidence I regard the dites hispidus, Foveotriletes parviretis, LaeviMurospora florida Zone to range from Neo- gatosporites sp., a small monolete, granulate comian to early Aptian in age. The precise form, and Rouseisporites reticulatus. These age of its lower limit in Queensland is subject types have doubtful stratigraphic significance to further discussion. At present it seems because of their infrequent occurrence. The reasonable to place it near the Jurassic-Cre- subzone contained also single specimens of taceous boundary. Cyathidites asper, Dictyotosporites filosus, Further remarks: The changes in the micro- Concavissimisporites penolaensis, and a small floral record between the Upper Jurassic and trilete spore characterized by a loosely enthe Aptian rank among the highest in impor- veloping (perinous) outer exine layer, which tance that are known in the Mesozoic of is more common in higher levels. None of Australia. The range chart prepared for the these species has been found beneath the present study (Table IV) shows the regular subzone; Dettmann (1963) recorded D. filosus occurrence of 52 spore and pollen species in and C. penolaensis from younger (Albian) Evans' unit J 6. For the Aptian, i.e. Evans' microfloras in southeastern Australia, and unit K lb-c, the count is at least 82 species, Dettmann & Playford (1969) indicated the Spec.Publs geol.Soe.Aust., 4: pp. 87-118, Pis 1-2, 1973.

98

D. B1

presence of D. filosus in the upper part of the Dictyotosporites speciosus Zone, the correlate of Evans' (1966a) unit K Id. Higher in the subzone the first specimens of Reticuloidosporites arcus were encountered, together with the first convincing specimens of Crybelosporites stylosus. The presence of this species near the lower limit of the subzone in Amerada Newlands No. 1 at 4,156 feet (1,266.8 m) is suspect, as Evans (1966d) thought that the sample was contaminated. It seems therefore, that two intervals can be recognized within the subzone, of which the upper one is distinguished by the occurrence of C. stylosus and R. arcus, and perhaps also Rouseisporites simplex, the earliest specimens of which were found in the interval of the subzone in Marchmont No. 1 and Whyenbirra No. 1 (Table III). The upper limit of the subzone is marked by the first appearance of Foraminisporis wonthaggiensis and Cicatricosisporites ludbrooki. Occurrence: Standard for the subzone is taken in BMR Roma No. 1 Scout Hole, Surat Basin, between core 40 at 406 feet (123.5 m) and core 37 at 377 feet (114.0 m ) . The subzone is in the Surat Basin associated with the upper part of the Orallo Formation, the Mooga Sandstone and the basal Kingull Member of the Bungil Formation (Fig. 2). One microflora from the upper Orallo Formation in AAO No. 1 (Roma) at 835 feet (254.5 m) was relatively poor in number of species, and did not contain Cicatricosisporites, but R. J. Paten (pers. comm.) of the Queensland Department of Mines in Brisbane, recovered C. australiensis from a sample of the same core. In the Eromanga Basin the subzone is associated with the Hooray Sandstone (Fig. 2). Towards the southeast the subzone was identified in Whyenbirra No. 1 in a sequence which is difficult to compare with that in the central and northern part of the basin, but which Graham (in press) regarded as the Hooray Sandstone (Fig. 4 ) . In the Maneroo Sheet area (Newlands No. 1) (Fig. 5) the subzone commences in the middle member of the Lower Hooray Sandstone; the lower part of the member seems still to be associated with unit J 5-6, judging by Poumot's (in Zolnai, 1965) examination of a microflora from 4,236 feet (1,290.8 m) in Mayneside No. 1. The upper limit of the subzone probably coincides

approximately with the top of the Lower Hooray Sandstone in Penrith No. 1 (Fig. 3) and Newlands No. 1. Towards the south there are no data concerning the subzone. In Orient No. 2 the upper member of the Lower Hooray Sandstone is associated with the Foraminisporis wonthaggiensis Subzone. The Cicatricosisporites australiensis Subzone may here be associated with the lower part of that member and with part, if not all, of the underlying middle member (Fig. 5). In the Longreach Sheet area, from near the base of the Hooray Sandstone in Saltern Creek No. 1 (Fig. 3) a microfioral assemblage was recovered, which contains Cicatricosisporites ludbrooki and Densoisporites velatus and is therefore regarded as part of the Foraminisporis wonthaggiensis Subzone (Table III). In the Injune Creek Group of the same well mid-Jurassic spores occur (Hodgson, in Mott & Associates, 1964a). This agrees with a generally accepted mid-Jurassic age for the group in the Great Artesian Basin (Evans, 1966a; Burger, 1968). The Cicatricosisporites australiensis Subzone is therefore not represented in this section. The same situation exists in Alice River No. 1, where an assemblage above the base of the Hooray Sandstone also represents the Foraminisporis wonthaggiensis Subzone. Exon et al (1966) noticed the lower part of the Hooray Sandstone missing from the sequence in outcrop in the Tambo Sheet area, east of the Enniskillen Anticline. This is most likely also the case in the Saltern Creek and Alice River wells. Age: There is little direct evidence for the age of the Cicatricosisporites australiensis Subzone. The earliest occurrence of C. australiensis is accompanied by the first appearance of at least 11 spore species, i.e. about 15% of the total number of species recorded at that level (Table IV). This point represents the largest numerical and percentage change in the Upper Jurassic-Lower Cretaceous sequence and is tentatively taken as the Jurassic-Cretaceous boundary in the Great Artesian Basin. The subzone is therefore at present regarded as of mainly early Neocomian (approximately Berriasian?) age. In Western Australia, Balme (1964) recorded equally important changes at the commencement of his Microcachryidites Assemblage, where C. australiensis is introduced. He thought, however, that these might have taken

SPORE ZONATION A N D HISTORY OF NEOCOMIAN 99 place in the uppermost Jurassic (late Titho- top of the Hooray Sandstone correlate in nian). Ooroonoo No. 1 (Fig. 6). In BMR Hughenden No. 1 both the Cicatricosisporites austraTHE Foraminisporis wonthaggiensis SUBZONE liensis and Foraminisporis wonthaggiensis Characteristics: The Foraminisporis wonthag- Subzones are missing. In the Maneroo Sheet area the subzone giensis Subzone is the middle of the three Subzones in the Murospora fiorida Zone. It appears to be entirely confined to the Upper Hooray Sandstone. In Orient No. 2 farther contains almost all of the species in the preceding subzone, but is distinguished from that south, the subzone includes also a considersubzone by the first appearance of F. won- able part of the upper member of the Lower thaggiensis and Cicatricosisporites ludbrooki. Hooray Sandstone. The upper limit of the The following species also seem to make their subzone lies definitely below the base of the first (sporadic) appearance within the sub- Rolling Downs Group in Innamincka No. 1, zone: Densoisporites velatus, Dictyotosporites Chandos No. 1, Cothalow No. 1, WOL No. speciosus, Januasporites spinulosus, Kuyli- 3 (Warbreccan), and Mayneside No. 1. sporites lunaris, and Kraeuselisporites majus. Age: The subzone is considered to be of NeoSpecies apparently not extending above the comian age. It is probably older than Barresubzone are a granulate monolete species, mian, as it lacks certain spore species which Aequitriradites hispidus, and Reticuloido- occur in younger intervals and which in other sporites arcus. This agrees with Dettmann's parts of the world are not known below the (1963) observations of restricted occurrence, Barremian. It is not possible to fix an accurate within the Stylosus Assemblage, of the latter lower age limit for the subzone. The presence two species. One specimen of Trilobosporites of Januasporites might suggest sp. which resembles T. apiverrucatus Couper, that it is not olderspinulosus Valanginian, as in 1958, was found in Saltern Creek No. 1. It also Canada species of thethan same genus from occurs infrequently in microfloras preceding the the Valanginian upwards, frequentlyoccur in abunMurospora fiorida Zone (unpublished infor- dance (Pocock, 1967). mation), and may be stratigraphically important. No specimens of the species were THE Foraminisporis asymmetricus SUBZONE found in the Cicatricosisporites australiensis Characteristics: The Foraminisporis asymSubzone. metricus Subzone is the highest of the three The subzone ends with the first appearance subzones which form the Murospora fiorida of F. asymmetricus and Pilosisporites notensis. Zone. It is distinguished from the previous Occurrence: The existence of the subzone was subzone by the first appearance of Pilosiestablished throughout the Surat and Ero- sporites notensis and F. asymmetricus. It conmanga Basins. Standard for the subzone is tains most of the species occurring in the taken in BMR Roma No. 1 Scout Hole in the other subzones, except for Aequitriradites his348-307 feet (106.1-93.6 m) interval (Fig. 2). pidus, Reticuloidosporites arcus, and a small In the Surat Basin the subzone is associated monolete form. The subzone includes the with the major part of the Kingull Member, earliest occurrence of Trilobosporites purveruBungil Formation. lentus and probably also Cyathidites puncIn the Eromanga Basin it is associated with tatus; it includes the last occurrence of the Hooray Sandstone, and was also found in Concavissimisporites sp. the Ronlow Beds. Towards the east it occurs Single specimens of the following species in BMR Mitchell No. 7, BMR Tambo No. were found within the subzone: Mineri5 and Alice River No. 1 (Fig. 2). In Alice sporites sp., Cooksonites variabilis, cf. DictyoRiver No. 1, Saltern Creek No. 1 and possibly phyllidites pectinataeformis, Pilosisporites also in Brookwood No. 1 the subzone is at parvispinosus, and Aequitriradites tilchaensis. least in part associated with the Lower Hooray The presence of P. parvispinosus and D. pecSandstone (Fig. 3). Farther to the west the tinataeformis is very unusual, as it represents subzone occurs in the Upper Hooray Sand- a considerable downwards extension of their stone of Yongala No. 1 and Orient No. 1 ranges, as known in eastern Australia. Dett(Fig. 5). It also occurs in the Upper Hooray mann (1963) recorded D. pectinataeformis of Whyenbirra No. 1, in the southeastern part from the Paradoxa Assemblage in the Otway of the basin (Fig. 4). In the northern part Basin, South Australia. In Queensland, the the subzone was recovered from near the species is known as a rare element in Evans' Spec.Publs geol.Soc.Aust, 4: pp. 87-118, Pis 1-2, 1973.

D. BURGER 100 (1966a) unit K 2 assemblages. P. parvi- covered (Fig. 6). Elsewhere in the northern spinosus is a commonly occurring species in part of the basin the Doncaster is not yet Evans' units K Id and K 2; Dettmann (loc. sufficiently known. A microflora from BMR cit.) recorded the species in the upper part Gilberton No. 2 at 25.9 m contained, among of her Speciosus Assemblage and Paradoxa others, Pilosisporites notensis and cf. MuroAssemblage. The presence of the two species spora florida. Cuttings from 700 feet (213 m) as low as the Foraminisporis asymmetricus in St Andrew's Bore (RN 14338) yielded an Subzone possibly indicates a hitherto unknown assemblage in which one specimen of M. florida was found. Farther south, in Brookdownwards extension of their ranges. Inaperturopollenites limbatus has until now wood No. 1, a microflora from 1,992 feet (Fig. not been found in assemblages younger than 3) contained cf. Crybelosporites stylosus, M. the Murospora florida Zone. Its full range in florida, and P. notensis. It is still too early to the Great Artesian Basin is not known, but evaluate these occurrences. The assemblages it may occur already prior to the zone in the might be younger than the Foraminisporis Surat Basin. Balme (1957, 1964) reported the asymmetricus Subzone; isolated occurrences species from his 'Microflora II-B' Micro- of M. florida and C. stylosus may well be due cachryidites Assemblage in Western Australia. to recycling by erosion of older sediments. Megaspores are extremely rare in the inter- The microfloras are provisionally assigned to val of the zone. A few specimens which are the subzone; the association of the subzone most probably to be referred to the genus with the Doncaster will be more extensively Minerisporites Potonie 1956, occurred in the investigated with forthcoming study of the Foraminisporis asymmetricus Subzone of Rolling Downs Group. Mitchell No. 11 (Table I). Age: At first sight the faunal evidence for Occurrence: The full extent of the subzone is the age of the subzone seems contradictory. still insufficiently known; the results of forth- The occurrence of the subzone in the Nullacoming work have to be awaited (p. 103). wurt and Minmi Members, Surat Basin, Standard for the subzone is for the time being suggests a ?Neocomian to early Aptian age, taken in BMR Roma No. 1 Scout Hole, Surat based on the faunas in these members. HowBasin, as the interval between core 26 at 219 ever, its inferred occurrence in the Doncaster feet (85.0 m) and core 16 at 185 feet (56.4 Member of Ooroonoo No. 1 suggests a late m). In the Surat Basin the subzone was Aptian age on account of the 'Roma' faunas identified in the Nullawurt and lower part found in outcrop of the member (Fig. 7). If of the Minmi Members of the Bungil Forma- the Doncaster is indeed consistently of late tion (Fig. 2). Farther west, in BMR Auga- Aptian age, then the Nullawurt and Minmi thella No. 3, the subzone appears to extend Members, being contemporaneous sequences slightly into the Doncaster Member of the in near-shore and partly nonmarine environments, should equally be regarded as late Wallumbilla Formation (Fig. 2). In the Eromanga Basin, in W.O.L. No. 3 Aptian in age. This assumption contradicts the (Warbreccan), Cothalow No. 1, Whyenbirra faunal evidence and for that reason cannot be No. 1, Mayneside No. 1 (see Poumot, in upheld. I prefer to adhere to Day's age estiZolnai, 1965), and very probably also Inna- mations for the Nullawurt and Minmi faunas mincka No. 1, the subzone was identified in and thus accept a ?Neocomian to early Aptian the Upper Hooray Sandstone (Figs 4, 5). The age for the Foraminisporis asymmetricus subzone also occurs in the Gilbert River Subzone. This implies that in the northern Eromanga Formation of BMR Hughenden No. 1 and BMR Gilberton No. 1 (Fig. 6). It may be Basin the Doncaster Member is in part locally absent in the Ronlow Beds (Fig. 3); of similar age. Here, palaeontological evipart of the rock sequence is missing in BMR dence does not exist south of the Hughenden Longreach No. 3, where the Doncaster rests and Richmond Sheet areas (Vine & Day, unconformably on the Ronlow Beds (Vine & 1965), except in Ooroonoo No. 1, where Belford (in McPhee, 1963) reported foraGalloway, 1969). In Ooroonoo No. 1 the subzone appears to minifera of presumably Lower Cretaceous age be associated with the Doncaster, as from the from the Doncaster at 2,298 feet (700.7 m). top of the underlying Hooray Sandstone cor- The member seems thus to be slightly diarelate a microflora from the preceding Fora- chronous. Indirectly, a Neocomian age for the lower minisporis wonthaggiensis Subzone was re-

SPORE ZONATION AND HISTORY OF NEOCOMIAN Ooroonoo No 1

101

Mitdwll No 7

Mitcholl Noll

Augathoito No 3

Roma No 1

Tom bo No 5

i

I

1 DONCASTER _

—-— Subzone of Foraminisporis asymmetricus

— HOORAY

8

CORRELATES

lithological boundary

——

_

M1M M 1 m 1IM IVI1

——

<™

^

Late Aptian "Roma" faunas

^ty

Early Aptian faunas

~~ NULLAWURT

^19 T Neocomian faunas

^KINGULL palynological boundary

Fig. 7. Stratigraphic position of the Foraminisporis asymmetricus Subzone (schematic representation). limit of the Foraminisporis asymmetricus Subzone is suggested by the presence of a few species which have been mentioned in the literature from Barremian and younger strata only. F. asymmetricus is known from the Barremian onwards in the USSR (Chlonova, 1969), in Britain (Kemp, 1970) and probably also in Canada (Pocock, 1962; Singh, 1964). Trilobosporites purverulentus is known in the USSR from Barremian or younger strata (Verbitskaya, 1962; Samoilovich et al., 1961; for detailed references see Dettmann & Playford, 1969). The available evidence, therefore, points to a late Neocomian (Barremian)—early Aptian age for the subzone. DINOFLAGELLATES AND OTHER MICROFOSSILS Marine sediments occurred only in association with the Foraminisporis asymmetricus Subzone. The microfloras recovered contained fair numbers of microplankton species. Most of the identified species have been described by Cookson & Eisenack (1958, 1960) from Australian Lower Cretaceous sediments. Evans (I966e) noticed the restricted ranges of some species in the marine Upper Jurassic to Lower Cretaceous sequence in the Papuan Basin, and used these to establish a suite of dinoflagellate zones. Those that appeared to be associated with the Murospora florida Zone are a lower Zone of Cannosphaeropsis mirabilis/ Scriniodinium attadalense, and an upper Zone of Scriniodinium attadalense / Dingodinium cerviculum, which according to Evans lacks C. mirabilis. The environmentally restricted occurrence of the marine fossils, shown in Table V, do not permit full comparison of Evans' zones

in Papua with the sequence in the Great Artesian Basin. However, the presence of cf. S. attadalense and C. mirabilis together with D. cerviculum in the Foraminisporis asymmetricus Subzone would indicate that his lower and upper zones are to a large degree overlapping. Study of new material from the Papuan Basin has confirmed this. It thus seems that the last occurrence of C. mirabilis closes an interval in the dinoflagellate sequence which is equivalent to Evans' lower and upper zones and marks the beginning of his Zone of Dingodinium cerviculum, which has been observed repeatedly in the Doncaster Member of the Great Artesian Basin. In many instances, lack of faunal evidence prevents dating the occurrence of various species in Australia. In the present case, however, Neocomian affinity of Chlamydophorella nyei, Canningia colliveri, Cribroperidinium edwardsi, Muderongia tetracantha, and Odontochitina operculata is suggested by their occurrence in the Foraminisporis asymmetricus Subzone. In Iehi No. 1, Papuan Basin, the earliest specimens of these species were found in late Neocomian sediments, so that Table V may actually represent approximately the true lower limits of the ranges of these species in Queensland. The Australian observations agree with Gocht's (1957, 1959) reports of O. operculata and Muderongia cf. M. tetracantha from uppermost Hauterivian and younger strata in Germany. Acritarchs and other microfossils appeared to be less sensitive environmentally; some of them occur almost throughout the Murospora florida Zone. However, specimens of Botryococcus spp., which many observers regard as a freshwater alga, have in no instance been recovered together with dinaflagellates; neither

Spec.Publs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

D. BURGER 102 do they occur in the marine Doncaster & Playford (1969). From the basal Cretaceous of southeastern Australia, Dettmann described Member. The problematic organism 'Gen. et sp. her Stylosus Assemblage (Dettmann & Playford's indet. A' Eisenack & Cookson has not been Crybelosporites stylosus Zone), with, among encountered in microfloras beneath the Muro- other species, C. australiensis, M. florida, and spora florida Zone, neither does it occur in C. stylosus. As we have seen, in Queensland the palynological sequence above the zone. C. stylosus does probably not appear prior to Eisenack & Cookson (1960) considered the the upper part of the Cicatricosisporites ausform as 'Aptian or older'; they reported it, traliensis Subzone. The species extends into among other localities, from the 'Blythesdale the Foraminisporis asymmetricus Subzone Group' of Santos Oodnadatta No. 1, Ero- and has also been observed in the succeeding manga Basin, South Australia, in the same unit K lb-c. In the Papuan Basin it has been found, although sporadically, as high as the interval (1,227-32 feet) from which Dettmann upper of the range of M. florida (unpub(1963) reported a microflora of her Speciosus lished limit information). Ingram (1968) reported Assemblage. The organism has not been the species the Eucla Basin together with observed in the Papuan area, and it does not Muderongia inmcwhaei, a form which in the seem to occur in the Cretaceous of Western Great Artesian Basin occurs in the ForaminiAustralia. sporis asymmetricus Subzone (Table V). In many instances only single specimens of the COMPARISON OF PALYNOLOGICAL species were found. ZONATIONS IN AUSTRALIA Dettmann's (1963) Speciosus Assemblage, Balme (1957) recorded the presence of Cicatricosisporites australiensis and Muro- Dettmann & Playford's (1969) Dictyotospora florida together in assemblages of his sporites speciosus Zone, contains D. speciosus 'Microflora II-B', with frequently occurring and lacks C. stylosus. The zone contains the Microcachryidites antarcticus. He found the first appearance of, among other species, ForaMicroflora in the Lower Cretaceous Perth minisporis asymmetricus. Also, M. florida Shale and ?Upper Jurassic Strathalbyn Sand- occurs in part of the zone. In the Great stone of the Perth Basin. In the Carnarvon Artesian Basin in South Australia, Dettmann Basin similar assemblages occurred in the (1963) recovered microfloras assigned to the (probably) Neocomian to lower Aptian Bird- lower part of the zone from the 'Roma Formation' in Cootabarlow No. 2 Bore at 1,376-7 rong Formation. In his review of Australian pre-Tertiary feet (493.5 m), containing M. florida, F. palynology, Balme (1964) distinguished in the asymmetricus, Cyathidites punctatus, and Western Australian region the Micro- Cooksonites variabilis; and also from the cachryidites and Hoegisporis Assemblages, 'Blythesdale Group' (or 'Cadna-Owie Formawhereby the Microcachryidites Assemblage tion'; see Dettmann & Playford, p. 187) in apparently includes the interval of 'Microflora Oodnadatta No. 1 at 1,227-32 feet (394 m), II-B'. The relationship of this assemblage to containing M. florida and Pilosisporites the Murospora florida Zone in Queensland is notensis (Dettmann, p. Ill, Table 6). Dettnot clear. The Hoegisporis Assemblage occurs mann did not record the presence of C. styin the (upper Albian to Cenomanian) Osborne losus in either assemblage. Formation in the Perth Basin. In Queensland, The stratigraphic position of these assemHoegisporis sp. has been found only in the blages is significant, as similar species assohigher interval of Evans' (1966a) spore units ciations are also known in Queensland from K 2, considerably beyond the Murospora the Foraminisporis asymmetricus Subzone. It florida Zone. The upper limit of the Micro- is further noted that Dettmann did not report cachryidites Assemblage must therefore lie F. asymmetricus, Cyathidites punctatus, or P. stratigraphically higher than the Murospora notensis in microfloras older than her Specioflorida Zone. sus Assemblage. Despite the presence of C. Lower Cretaceous sediments, rich in pollen, stylosus, the Foraminisporis asymmetricus are known in the whole of the eastern Aus- Subzone may thus be equivalent to the lower tralian-Papuan region. Microfloral sequences part of the Dictyotosporites speciosus Zone, from the Gippsland, Otway and Great or, more specifically, the lower part of the Artesian Basins have been studied by Dett- Cyclosporites hughesi Subzone. If this is mann (1963), Evans (1966<z), and Dettmann correct, then the Crybelosporites stylosus

SPORE ZONATION AND HISTORY OF NEOCOMIAN

Zone may be equivalent to the upper part of the Cicatricosisporites australiensis Subzone and the Foraminisporis wonthaggiensis Subzone in Queensland (Fig. 8). Dettmann & Playford (1969) regarded the Crybelosporites stylosus Zone as not younger than Valanginian. Dettmann (1963) did not record the nominate species from the marine 'Roma Formation' (the Doncaster Member correlate), but in the Surat Basin, Queensland, the nominate species also occurs in assemblages younger than unit Kla (Table I), and even in the Aptian Doncaster Member (unpublished data). This apparent discrepancy may partly be due to lack of faunal control; also, the possibility of recycling of specimens into the Aptian of Queensland should be taken into account. On present knowledge, however, C. stylosus appears to be a poor zone index species in Queensland and Papua. The concept of Evans' unit K la has led to identical results as those obtained by regional fieldwork, and thus promises to yield more reliable data with future palynostratigraphic studies in those regions. Dettmann (1963) and Evans (1966a) regarded the first appearance of C. australiensis as approximately coinciding with the JurassicCretaceous boundary. From Balme's (1964) observations on the oldest occurrence of this species in Western Australia, Dettmann & Playford (1969) considered an uppermost

BALMEJ957

BALME, 1964

103

Jurassic age for part of the Crybelosporites stylosus Zone possible. If C. australiensis indeed occurs in the uppermost Jurassic in eastern Australia, then the sequence from the point of its first occurrence up to the incoming of C. stylosus represents (entirely or in part) the closing portion of the Jurassic. If the first appearance of C. stylosus has been accurately determined in the pollen sequence of the Great Artesian Basin, Queensland, then the Crybelosporites stylosus Zone may be entirely of Cretaceous (Neocomian) age, according to Dettmann's initial views. CONCLUSIONS Study of the spore-pollen history in the Lower Cretaceous of the Great Artesian Basin, Queensland, which started a decade ago, has been progressing with rapidly increasing knowledge of the geology of the basin. Study of the basal Cretaceous in particular was intensified in recent years from the wealth of material that became available. Various aspects in the field of pollen stratigraphy were discussed in the previous pages. In the Neocomian to early Aptian of the Great Artesian Basin, three intervals were distinguished in the palynological sequence, which proved to be very useful for correlation of sediments in central and marginal areas of the Surat and Eromanga Basins, where other

DETTMANN a PLAYFORD, 1969 EVANS, 1966 BURGER (THIS

PAPER)

INTERN. STAGES

H0EGISPORIS Kid Klb-c

APTIAN

Cyclosporites hughesi Foraminisporis asymmefricus

MICROFLORA n-B

Kla CRYBELOSPORITES STYLOSUS

p

MUROSPORA FLORIDA

MICROCACHRYDITES

DICTYOTOSPORITES SPECIOSUS

ALBIAN Crybelosporites striatus

Foraminisporis wonthaggiensis

NEOCOMIAN

Cicatricosisporites australiensis UPPER JURASSIC

Fig. 8.

Correlation of palynological zonations in Australia.

Spec.PubIs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

104

D. BURGER

Subzone of

Subzone of

Subzone of

Cicatricosisporites australiensis

Foraminisporis wonthaggiensis

Foraminisporis asymmetricus

O 0

200 200

400

400

MILES

600

KILOMETRES

ISOPACHS OF ROCK COLUMNS ASSOCIATED WITH EACH OF THE SUBZONES (TENTATIVE Numbers

indicate

thickness in metres

Edge of Great Artesian Basin

•

RECONSTRUCTION) Control sections

Extent of Doncaster deposition In the l a t e Neocomian Extent of Doncaster deposition in the early A p t i o n Isopachs taken at 2 5 m intervals Numbers indicate estimated thickness (in metres) for selected bore sections Palaeolatitudes derived from Day (1969)

ISOPACHS OF STRATA ASSOCIATED WITH THE ZONE OF MUROSPORA FLORIDA

Fig. 9.

^

^

Marine influence outside area of Doncaster sedimentation Possible total extension of sea in the early A p t i a n Numbers are isopachs of the Doncaster in the interval of Subzone

TOTAL E X T E N T OF THE SEA IN THE SUBZONE OF FORAMINISPORIS ASYMMETRICUS

Palaeogeographic and isopach maps for the Neocomian-early Aptian in the Great Artesian Basin, Queensland.

SPORE ZONATION AND HISTORY OF NEOCOMIAN palaeontological evidence did not exist. These intervals, in stratigraphic order, the Cicatricosisporites australiensis Subzone, the Foraminisporis wonthaggiensis Subzone and the Foraminisporis asymmetricus Subzone, together form the Murospora fiorida Zone. Spore intervals equivalent to the zone have been reported in Lower Cretaceous sequences throughout Australia. On the basis of the evidence presented, an attempt is made to sketch the geological history of the Surat and Eromanga Basins in the Neocomian-early Aptian from a palynological point of view. Two major periods of sedimentation are distinguished. During the first period, of Neocomian age, fluviatile and lacustrine deposition was characteristic for most of the area. An early phase occurred with accumulation of the Mooga Sandstone in the Surat Basin from south-flowing streams (Exon et al., 1967) and contemporaneous deposition of sands (upper member of Lower Hooray Sandstone) in the central Eromanga Basin, presumably from the west (Vine et al., 1965). In the northeast, close to areas of erosion or non-deposition, quietly, southerly flowing streams accumulated the fine-grained arenaceous and argillaceous beds of the Hooray Sandstone in the TamboAugathella region (Exon et al., 1966). Isopachs of strata associated with this phase

105

(Fig. 9A) show the existence of two major areas of deposition, limited towards the northeast in the Hughenden and Longreach areas, and towards the east near Roma and Mitchell. The areas are separated by what are referred to as the Nebine and Eulo Ridges. A subsequent phase was characterized by predominantly fine-grained sediments of the Bungil Formation in the Surat Basin and the paralic silts and muds of the Upper Hooray Sandstone in the Maneroo region. Isopachs for this phase (Fig. 9B) show the thickest accumulation in the south, west of the Eulo Ridge, where deposition of the Lower Hooray Sandstone did not cease until a later stage. In the second major period, of upper Neocomian to early Aptian age, large areas in the north and east were flooded by the sea. Isopachs of strata associated with that period are shown in Figure 9C. Field evidence from the central Eromanga Basin suggests that sediments developed in fluviatile to deltaic environments (Gregory et al., 1967). Successive limits of marine Doncaster Member deposition (Fig. 9E) indicate that the open ocean advanced in southern to southeastern directions. Towards South Australia high areas may have persisted, considering that at the location of Innamincka No. 1 deposition of nonmarine Hooray Sandstone continued without interruption throughout the period. LOWERMOST ROLLING DOWNS GROUP (Doncaster)

( unit Kl b - c )

Minmi MUROSPORA

2

o

FLORIDA

(unit KI a )

Foraminisporis Nullawurt

asymmetricus Foraminisporis wonthaggiensis

Kingull

Cicatricosisporites australiensis

MOOGA ORALLO

(units

GUBBERAMUNDA

J ? 5-6) BLANT

PALYNOLOGICAL ZONATION E R O M A N G A

BASIN

SURAT

BASIN

Fig. 10. Palynological correlation of Upper Mesozoic rock formations in the Surat and Eromanga Basins. Spec.Publs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

106

D. BURGER

The ocean may have had simultaneous access to the Surat Basin from the east. This is suggested by the configuration of isopachs and by the presence of marine strata in the Bungil Formation in the Roma-Mitchell area, in the uppermost Hooray Sandstone of Whyenbirra No. 1, and even in the Eulo area (Burger, in Senior et al., 1969), in each case prior to Doncaster deposition. Day (1969) suggested that a corridor may have existed to the east, from the occurrence of similar forms of Meleagrinella in the Surat and Maryborough Basins. Soon after the close of this period the transgression reached its peak. The Doncaster Member and equivalent marine argillaceous strata occur throughout the Great Artesian Basin as far south as northern New South Wales and towards the west also in the Northern Territory and northern South Australia (Day, 1969). Palynological correlation of rock formations associated with the Murospora florida Zone is given in Figure 10. The climate during the Neocomian-early Aptian may have been cool to temperate (Dettmann & Playford, 1969). Day (1969) mentioned results of oxygen isotope analyses on Lower Cretaceous Peratobelus from South

Australia (Lake Eyre District), which point to cool palaeotemperatures. He regarded the 'Roma' fauna as a cool-water fauna because of its links with equivalent boreal faunas and the absence of temperate 'Tethys' genera. So far, the study of Mesozoic spores and pollen grains affords no clear intimation of the climate.

AMERADA PETROLEUM CORPORATION OF AUSTRALIA

BUSNARDO, R., LE HEGARAT, G., & MAGNE, J.,

ACKNOWLEDGMENTS Various colleagues in the Bureau of Mineral Resources have given considerable aid during preparation of the present study. I wish to thank in particular Messrs R. R. Vine, B. R. Senior, H. F. Doutch and J. Smart for instructive discussions on results of progressing (unpublished) field work and various problems on the stratigraphy in the Eromanga and Carpentaria Basins. Dr N. J. de Jersey of the Geological Survey of Queensland gave valuable comments on the manuscript, for which I am very grateful. I also wish to express my gratitude to Dr J. M. Dickins and Dr D. J. Belford for their valuable comments while reading the manuscript. The Director of the Bureau of Mineral Resources very kindly permitted publication of the manuscript in this volume.

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, INGRAM, J. A., THOMAS, B. M., & SENIOR, DANIELE, 1969: The geology of the

Quilpie, Charleville, Toompine, Wyandra, Eulo and Cunnamulla 1:250,000 Sheet areas, Queensland. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1969/13 [unpublished]. SENIOR, DANIELE, {in press): Thargomindah, Queensland 1:250,000 Geological Series. Explan. Notes Bur. Miner. Resour. Geol. Geophys. Aust., SG/54-16. SINGH, C., 1964: Microflora of the Lower Cretaceous Mannville Group, east-central Alberta. Bull. Res. Coun. Alberta, 15, pp. 1-238. TEICHERT, C., 1958: Some biostratigraphical concepts. Bull. geol. Soc. Am., 69, pp. 99-120. THOMAS, B. M., & REISER, R . F., 1968: T h e

geology of the Surat 1:250,000 Sheet area. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1968/56 [unpublished]. TWIST, R. F., 1962: Well Completion Report, pletion Report [unpublished]. Smart Oil Orient No. 2, Queensland, of MOTT, W. D., & ASSOCIATES, 1964A: Well ComL. H. Smart Oil Exploration Co. Ltd. [unpetion Report Saltern Creek No. 1 (Longpublished]. reach Oil Limited) [unpublished]. UNION OIL DEVELOPMENT CORPORATION, KERN , , 1964b: Marchmont No. 1 Well COUNTY LAND COMPANY, & AUSTRALIAN OIL Completion Report (Longreach Oil Limited) & GAS CORPORATION LTD., 1964: U . K . A . [unpublished]. Cabawin No. 1, Queensland. Bur. Miner. Resour. Aust. Petrol. Search Subs. Acts, 43. NORRIS, G., 1969: Miospores from the Purbeck Beds and marine Upper Jurassic of southern VEEVERS, J. J., & WELLS, A . T., 1961: T h e geology of the Canning Basin, Western AustraEngland. Palaeontology, 12 ( 4 ) , pp. 574-620. lia. Bull. Bur. Miner. Resour. Geol. GeoPEMBERTON, R. L., 1963: Well Completion Re60. port Exoil Brookwood No. 1 Well [unpub- VINE,phys.R . Aust., R., 1964a: Mackunda, Queensland lished]. 1:250,000 Geological Series. Explan. Notes POCOCK, S. A. J., 1962: Microfloral analysis and Bur. Miner. Resour. Geol. Geophys. Aust., age determination of strata at the JurassicSF/54-11. Cretaceous boundary in the western Canada , 1964b: Julia Creek, Queensland 1:250,000 plains. Palaeontographica B, 111, pp. 1-95. Geological Series. Explan. Notes Bur. Miner. Resour. Geol. Geophys. Aust., , 1967: The Jurassic-Cretaceous boundary SF/54-3. in northern Canada. Rev. Palaeobot. Paly, 1966: Recent geological mapping in the nol., 5, pp. 129-136. northern Eromanga Basin, Queensland. REYNOLDS, M. A., 1960: Mesozoic and younger APE A J. 1966, pp. 110-115. sediments of the Gilberton and Georgetown , CASEY, D . J., & JOHNSON, N . E. A., 1964: 4-mile Sheet areas, Queensland. Rec. Bur. Progress Report, 1963, on the geology of Miner. Resour. Geol. Geophys. Aust., part of the north-eastern Eromanga Basin, 1960/68 [unpublished]. Queensland. Rec. Bur. Miner Resour. Geol. RYAN, J. C., 1961: Innamincka No. 1 Well, South Geophys. Aust., 1964/39 [unpublished]. Australia. Bur. Miner. Resour. Aust. Petrol. , & DAY, R. W., 1965: Nomenclature of Search Subsidy Acts Publ., 9. the Rolling Downs Group, northern Eromanga Basin, Queensland. Qd Govt Min. SENIOR, B. R., 1969: Jundah, Queensland J., 66 (767), pp. 417-421. 1:250,000 Geological Series. Explan. Notes , & GALLOWAY, M . C., 1969: Shallow Bur. Miner. Resour. Geol. Geophys. Aust., stratigraphic drilling, northern Eromanga SG/54-4. Basin, 1963-64. Rec. Bur. Miner. Resour. , GALLOWAY, M . C., INGRAM, J. A., & Geol. Geophys. Aust., 1969/20 [unpubSENIOR, DANIELE, 1968: The geology of the lished]. Barrolka, Eromanga, Durham Downs, , & JAUNCEY, W., 1962: Explanatory Notes, Thargomindah, Tickalara and Bulloo Julia Creek Sheet, Queensland. Rec. Bur. 1:250,000 Sheet areas, Queensland. Rec. Miner. Resour. Geol. Geophys. Aust., Bur. Miner. Resour. Geol. Geophys. Aust., 1962/81 [unpublished]. 1968/35 [unpublished].

MINES ADMINISTRATION PTY LTD., 1963: Penrith No. 1 Well, A P 8 6 P , Queensland, Com-

SPORE ZONATION AND HISTORY OF NEOCOMIAN

109

, JAUNCEY, W . , CASEY, D . J . & GALLOWAY,

WHITEHOUSE, F . W., 1955: T h e geology of the

M. C., 1965: Geology of the LongreachJericho-Lake Buchanan area, Queensland. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1965/245 [unpublished]. W H I T E , MARY E . , 1 9 6 3 : Plant fossil collections from Hughenden area, Great Artesian Basin. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1964/64 [unpublished]. , 1967: Report on 1966 collections of plant fossils from the Surat Basin, southwest Eromanga Basin, Delamere, Northern Territory: and Proserpine district of Queensland. Rec. Bur. Miner. Resour. Geol. Geophys., 1967/78 [unpublished].

Queensland portion of the Great Australian Artesian Basin. Artesian Water Supplies in Queensland 1954; Append. G. Dep. Coord. Gen. Publ. Works Pari. Pap. A, 56-1955 {Brisbane). WOOLLEY, J. B., 1941: Geological report on the area northeast of Tambo. Shell (Queensland) Development Pty Ltd. Rep. 5 [unpublished]. ZOLNAI, G., 1965: Australian Aquitaine Petroleum Pty Ltd. Mayneside No. 1, AP 86 P, Queensland, Well Completion Report [unpublished].

D. Burger, Bureau of Mineral Resources, Geology and Geophysics, P.O. Box 378, Canberra City, A.C.T. 2601.

Spec.Publs geol.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

D. BURGER TABLE I

110

Stratigraphic occurrence of tnicrofossils in the Surat Basin

g!

? Probable occurrence of species MICROFOSSIL

SPECIES

SPORES AND POLLEN GRAINS AEQUITR1RADITE8 HISPIDUS A. SPINULOSUS A. VERRUCOSUS AUSPORITES GRANDIS A. SI Ml LIS ANNULJSPORA FOLLICULOSA ARAUCAR LA CITES AUSTRALIS BACULATISPORITES COMAUMENSIS BIRETISPORITES cf. B. POTONIAEI B. SPECTABIUS CALLIALASPORITES DAMPIERI C. TRILOBATU8 CERATOBFORITES EQUALIS CICATRICOSISPORITKS AUSTRALIENSIS C. HUGHE8I C. LUDBROOKI C. sp. CINGUTRILETES CLAVU8 CLASSOPOLLIS spp. CONCAVISPOHITES spp. CONCAVISSIMISPORITES PENOLAENSIS C. Bp. CONCENTRISPORITES HALLEI CONTIGNISPORITES COOKSONH C FORNICATUS C. MULTIMURATUS C. spp. Indot. COOKSONITES VARIABILIS CORONATISPORA PERFORATA COUPERISPORITES TABULATU8 CRTBE LOSPORITES STYLOSUS CYATHIDITE8 AUSTRALIS C. MINOR CYCLOSPORITES HUGHES! DENSOISPORITES VELA TITS IXCTYOPHYLUDITES CRENATU8 D. MORTONI

D. PECTINATAEFORMIS OtCTYOTOSPORITES COMPLEX O. FILOSUS D. SPECIOSU8 FORAMINISPORIS ASYMMETRICUS P. DAILY1 F. WONTHAGGIENSB POVEOSFORITES CANALIS FOVEOTRILETES PARVTRETIS GLEICHE NHDITES CIRCINIDTTE8 INAPERTUROPOLLENITES TURBATUS ISCHYOSPORITES PUNCTATUS JANUASPORITES SPINULOSUS KLUKISPORITES SCABERIS KRAEUSELISPORITES LINEARIS K. MAJUS AUYLISPORITES LUNARIS LAEVIGATOSPORITES sp. LEPTOLEPIDITES MAJOR L VERRUCATUS LYCOPODIACIDITE8 AMBIFOVEOLATUS LYCOPODIUMSPORITE8 AUSTROCLAVATTDITES L. CIRCOLUMENUS L. EMINULU8 L. FACETUS L. NODOSU8 L. RETICULUMSPORITE8 L ROSEWOODENSIS L. SEMIMURUS MATONISPORITE8 COOKSONH MICROCACHKYIDITES ANTARCTICU8 MINERI8PORITES sp. MONOSULCITES MINIMUS MUR06P0RA FLORIDA NEORAI8TRICKIA TRUNCATA NEVE3ISPORITE8 VALLATUS 06MUNDACIDITES WELLMANH PERINATE INAPERTURATE POLLEN TYIPE PERINATE TRILETE SPORE PI IjOSISPORITES NOTEN8IS P. PARVISPINOSUS PODOCARPTOTES ELLIPTICUS P. MULTE8EMUS PC J.YCINGULATISPORITES DENSATUS RETICULATBPORITES PUDENS RETICULOIDOSPORITES ARCUS ROUSEISPORITES RETICULATUS S. SIMPLEX SESTR06P0RITES PSEUDOALVEOLATUS STEHE1SPORITES ANTTQUASPORITES TRIUTE8 cf. T. TUBERCULIFORMIS THJ ijOBOSPORITES ANTIQUUS TRILOBOfl PORXTES PURVERULENTUS TSI'f.iAEPOLLEN!TES 8EGMENTATUS VEUJSPORITES TRIQUETRUS VI raEISPORITES PALLIDU8 CAMNDKHA COLLIVERI CHLAMYDOPHORSLLA NTEI CHIBBOPERIDINIUM EDWARDS! CYCLONEPHEUUM DISTINCTUM DJNQODCNIUM CE3VICULUM GONIAULACY8TA HELICOIDEA MWDERONGIA MCWHAEI M. TETHACANTHA ODONTOCHSTINA OPERCULATA OUGOSPHAEBIDIUM ANTHOPHORUM TENUA sp. POTRYOOOCCUS spp. •CEN. ET SP. INDET. A* EU. * Cooks. I960 PTERG6PERMOP8I8 AUSTRALIENSIS SCK'ZOTPORIS RETICULATUS MlCRHYSTHIDniM spp.

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SPORE Z O N A T I O N A N D HISTORY OF N E O C O M I A N TABLE I I

Stratigraphic occurrence of microfossils in the central Eromanga Basin ROCK

UNIT

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Spec.Publs geol.Soc.Aust, 4: pp. 87-118, Pis 1-2, 1973.

MESOZOIC UNDIFF. Tiorida

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D. B U R G E R TABLE III

Stratigraphic occurrence of microfossils in the northern and eastern Eromanga Basin

Positive identification of species

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Probable occurrence of species MICROFOSSIL

SPECIES

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SWtern Creek No. 1 e. 3, 1664' Tambo No. 5 e., 77-11" ' Broolcwood No. 1 e. 5, 2110' Mitchell No. 7 c. 10, 180'8" Alice River No. I c. 2, 860-78' Whyenblrra No. 1 ewe, 1600' Whyenbirra No. 1 swc, 1400'

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Cicatricosisporites australiensis

ZONATION

Jericho No. 1 c. 1, 85'11" Longreach No. 1 c., 153' GUberton No. 2 c. 7, 34.43M Hughenden No. 1 c. 4, 202' GUberton No. 2 c. 4, 25.87m

H

UNIT

SAMPLE SPECIFICATION

ROCK

PALYNOLOGICAL

• •

• ? ? • • •

t • • ? • T

• • •

•

• • •

••

• •

SPORE ZONATION AND HISTORY OF NEOCOMIAN

113

TABLE I V

General distribution of spores and pollen grains in the Upper Mesozoic of the Great Artesian Basin Presence based on actual observation

ZONE

Presence induced from observations of species in other spore intervals Possible presence or slight extension of range of species on the basis of doubtful specimens outside actual range

AEQUITRIRADITES SPINULOSUS (Cooks. & Dettm., 1958) Cooks. & Dettm., 1961 A. VERRUCOSUS (Cooks. & Dettm., 1958) Cooks. & Dettm., 1961 ALISPORITES GRANDIS (Cookson, 1953) Dettmann, 1963 A. SI MI LIS (Balme, 1957) Dettmann, 1963 ANNULISPORA FOLLICULOSA (Rogalska. 1954) De Jersey, 1959 ARAUCARIACITES AUSTRALIS Cookson, 1947 BACULATISPORITES COMAUMENSIS (Cookson, 1953) Potonie, 1956 BIRETISPORITES SPECTABILIS Dettmann, 1963 CALLIALASPORITES DAMPIERI (Balme, 1957) Dev, 1959 C. TRILOBATUS (Balme, 1957) Dev, 1959 CERATOSPORITES EQUALIS Cooks. & Dettm., 1958 CINGUTRILETES CLAVUS (Balme, 1957) Dettmann, 1963 CLASSOPOLLIS spp. CONCAVISPORITES spp. CONCENTRISPORITES HALLEI (Nilsson, 1958) Wall, 1965 CONTIGNISPORITES COOKSONII (Balme, 1957) Dettmann, 1963 C. FORNICATUS Dettmann, 1963 CORONATISPORA PERFORATA Dettmann, 1963 CYATHIDITES AUSTRALIS Couper, 1953 C. MINOR Couper, 1953 DICTYOPHYLLIDITES CRENATUS Dettmann, 1963 D. MORTONI (De Jersey, 1959) Playford & Dettm., 1965 FORAMINISPORIS DAILYI (Cooks. & Dettm., 1958) Dettmann, 1963 FOVEOSPORITES C ANA LIS Balme, 1957 GLEICHENnDITES CIRCINIDITES (Cookson, 1953) Dettmann, 1963 INAPERTUROPOLLENITES TURBATUS Balme, 1957 ISCHYOSPORITES PUNCTATUS Cooks. & Dettm., 1958 KLUKISPORITES SCABERIS (Cooks. & Dettm., 1958) Dettmann, 1963 LEPTOLEPIDITES VERRUCATUS Couper, 1953 LYCOPODIACIDITES AMBIFOVEOLATUS Brenner, 1963 LYCOPODIUMSPORITES AUSTROCLAVATIDITES (Cookson, 1953) Potonie, 1956 L. CIRCOLUMENUS Cooks. & Dettm., 1958 L. EMINULUS Dettmann, 1963 L. NODOSUS Dettmann, 1963 L. RETICULUMSPORITES (Rouse, 1959) Dettmann, 1963 L. ROSEWOODENSIS (De Jersey, 1959) De Jersey, 1963 MICROCACHRYIDITES ANTARCTICUS Cookson, 1947 MONOSULCITES MINIMUS Cookson, 1947 NEORAISTRICKIA TRUNCATA (Cookson, 1953) Potonie, 1956 OSMUNDACIDITES WELLMANH Couper, 1953 PODOCARPIDITES ELLIPTICUS Cookson, 1947 P. MULTESIMUS (Bolchovitina, 1956) Pocock, 1962 POLYCINGULAT1SPORITES DENSATUS (De Jersey, 1959) Playford & Dettm., 1965 SESTROSPORITES PSEUDOALVEOLATUS (Couper, 1958) Dettmann, 1963 STAPLINISPORITES CAMINUS (Balme, 1957) Pocock, 1962 STER EISPORITES ANTIQUASPORITES (Wilson & Webster, 1946) Dettmann,1983 TRILOBOSPORITES ANTIQUUS Reiser & Williams, 1969 TSUGAEPOLLENITES SEGMENTATUS (Balme, 1957) Dettmann, 1963 VITREISPORITES PALLIDUS (Reissinger, 1938; 1950) Nilsson, 1958 LYCOPODIUMSPORITES SEMIMURUS (Danze-Corsin & Lav.. 1963) Reis. & Will., 1969 PERINATE INAPERTURATE POLLEN TYPE CONCAVISSIMISPORITES sp. MUROSPORA FLORIDA (Balme, 1957) Pocock, 1961 NEVESISPORITES VALLATUS De Jersey & Paten, 1964 TRILOBOSPORITES sp. INAPERTUROPOLLENITES LIMBATUS Balme, 1957 ISCHYOSPORITES MARBURGENSIS De Jersey, 1963 CONTIGNISPORITES MULTIMURATUS Dettmann, 1963 DICTYOTOSPORITES COMPLEX Cookson & Dettmann, 1958 LEPTOLEPIDITES MAJOR Couper, 1958 MATON1SPORITES COOKSONn Dettmann, 1963 BIRETISPORITES cf. B. POTON1AEI Delcourt & Sprumont., 1955 CYATHIDITES CONCAVUS (Bolchovitina, 1953) Dettmann, 1963 RETICULATISPORITES PUDENS Balme, 1957 TRILITES cf. T. TUBERCULIFORMIS Cookson. 1947 VELOSPORITES TRIQUETRUS (Lantz, 1958) Dettmann, 1963 CYATHIDITES ASPER (Bolchovitina, 1953) Dettmann, 1963 AEQUITRIRADITES fflSPIDUS Dettmann & Playford, 1968 GRANULATE MONOLETE SPECIES CONCAVISSIMISPORITES PENOLAENSIS Dettmann, 1963 CICATRICOSISPORITES AUSTRALIENS^ (Cookson, 1953) Potonie, 1956 C. HUGHESI Dettmann, 1963 C. sp. COUPERISPORITES TABULATUS Dettmann, 1963 CYCLOSPORITES HUGHESI (Cooks. & Dettm., 1958) Cooks. & Dettm., 1959a FOVEOTRILETES PARVIRETIS (Balme, 1957) Dettmann, 1963 LAEVIGATOSPORITES sp. ROUSEISPORITES RETICULATUS Pocock, 1962 RETICULOIDOSPORITES ARCUS (Balme, 1957) Dettmann, 1963 CRYBELOSPORITES STYLOSUS Dettmann, 1963 DICTYOTOSPORITES FILOSUS Dettmann, 1963 PERINATE TRILETE SPORE ROUSEISPORITES SIMPLEX (Cooks. & Dettm., 1958) Dettmann, 1963 LYCOPODIUMSPORITES FACETUS Dettmann, 1963 CICATRICOSISPORITES LUDBROOKI Dettmann, 1963 KRAEUSELISPORITES LINEARIS (Cooks. & Dettm., 1958) Dettmann, 1963 DENSOISPORITES VELATUS Weyland & Kriegfer, 1953 emend. Krasnova, 1961 DICTYOTOSPORITES SPECIOSUS Cooks. & Dettm., 1958 FORAMINISPORIS WONTHAGGIENSXS (Cooks. & Dettm., 1958) Dettmann, 1963 JANUASPORITES SPINULOSUS Dettmann, 1963 KRAEUSELISPORITES MAJUS (Cooks. & Dettm., 1958) Dettmann, 1963 KUYLISPORITES LUNARIS Cooks. & Dettm., 1958 MINERISPORITES sp. COOKSONITES VARIABILIS Pocock, 1962 TRILOBOSPORITES PURVERULENTUS (Verbitskaya, 1962) Dettmann, 1963 CYATHIDITES PUNCTATUS (Delcourt & Sprumont, 1955) Delcourt et al., 1963 FORAMINISPORIS ASYMMETRICUS (Cooks. & Dettm., 1958) Dettmann, 1963 PILOSISPORITES NOTENSIS Cooks. & Dettm., 1958 DICTYOPHYLLIDITES PECTINATAEFORMIS (Bolchovitina, 1953) Dettmann, 1963 AEQUITRIRADITES TILCHAENSIS (Cooks. & Dettm., 1958) Cooks. & Dettm., 1961 PILOSISPORITES PARVISPINOSUS Dettmann, 1963

Spec.Publs geal.Soc.Aust., 4: pp. 87-118, Pis 1-2, 1973.

MUROSPORA

FLORIDA

Cicatricosisporites

Foraminisporis

Foraminisporis

australiensis

wonthaggiensis

asymmetricus

114

D.

BURGER

TABLE V

General distribution of dinoflagellates and other microfossils in the Upper Mesozoic of the Great Artesian Basin Presence based on actual observation •••••

Presence induced from observations of species in other spore intervals

y

Possible presence or slight extension of range of species on the basis

ZONE

MUROSPORA

FLORIDA

Cicatricosisporites

Foraminisporis

Foraminisporis

austrahensis

wonthaggtensis

asymmetricus

Subzone

BOTRYOCOCCU8 spp. "GEN. ET SP. INDET. FORMA A* Eis. ft Cooks., 1060 SCHIZOSPORIS RETICULATUS Cooks, ft Dettm., 1950b S. PARVUS Cooks, ft Dettm., 1050b MICRHYSTRIDIUM SDD PTEROSPERMOPS1S AUSTRALXENSIS Deflandre ft Cookson, 1055

1

I f !

1

•

*

| =

f i l l

III

(

1

1

-

| O T H E R S ||

OLIGOSPHAERIDIUM ANTHOPHORUM (Cooks. ft Els., 1058) Davey et al., 1966 GONIAULACYSTA HELICOIDEA (Els. ft Cooks., 1060) Davey et al., 1066 DEFLANDREA sp. cf. PALAEOSTOMOCYSTIS sp. CRIBROPERIDINIUM EDWARDSI (Cooks. ft Els., 1058) Davey, 1060 DINGODINIUM CERVICULUM Cooks. & Eis., 1058 CYCLONSPHELIUM DISTINCTUM Defl. ft Cooks., 1055 CANNINGIA COLLTVERI Cooks. ft Eis., 1060 CHLAMYDOPHORELLA NYEI Cooks, ft Eis.. 1058 ODONTOCHITINA OPERCULATA (Wetzel, 1033) Defl. ft Cooks., 1055 SCRINIOMNIUM ATTADALENSfi (Cooks. ft Eis., 1058) Downie & Sarjeant, 1066 CANNOSPHAEROPSIS M1RABILIS Cooks. ft Eis., 1058 MUDERONGIA MCWHAEI Cooks. ft Eis., 1058 M. TETRACANTHA (Gocht, 1057) Albertl, 1061 TENUA sp.

DINOFLAGELLATES

of doubtful specimens outside actual range

SPORE ZONATION AND HISTORY OF NEOCOMIAN APPENDIX A Specifications of oil exploration wells depicted in Figs 2-6 Well

Latitude (South)

Longitude Year (East) Completed

1:250,000 Sheet Area

Reference

F.D. Alice 23°37'8" 145°19'15" River No. 1 Exoil Brook- 22°28'55" 144°19'58" wood No. 1 U.K.A. Caba- 27°29'46" 150°11'22" win No. 1

1963 Longreach 1962 Muttaburra 1961 Dalby

Hare & Associates, 1963 Pemberton, 1963 Union Oil Corporation et al., 1964

P.S. Cothalow 25°43'47" 144 23'41" No. 1

1961 Adavale

Lewis, 1961

Alliance 25° 50'14" 143°19'36" Chandos No. 1 D.F.S. Inna- 21°29'2\" 140°55'7" mincka No. 1

1966 Windorah 1959 Innamincka*

Laing & Benedek, 1966 Ryan, 1961

L.O.L. March- 23°10'15" 144°44'21" mont No. 1 A.A.P. Mayne- 23°35'23" i42°3rir side No. 1

1964 Longreach 1964 Maneroo

Mott & Associates, 19646 Zolnai, 1965

A.P.C. Newlands No. 1

1966 Maneroo

Amerada Petroleum Corporation of Australia Ltd., 1966

1960 Brighton Dns

McPhee, 1963

0

23°52'33" 142°57 46 /

/r

Conorada 23°10'50" 141°33'09" Ooroonoo No. 1 Smart Oil 27°40'30" 143°11'45" Orient No. 2 A.A.O. Pen- 23°10'20' 143°38'30" rith No. 1 A.A.O. Roma 26°34'09" 148°51'10" No. 1 L.O.L. Saltern 23°20'54" 144°56'24" Creek No. 1 W.O.L. No. 2 24°25'30" 142°47'00" (Warbreccan) /

1962 Thargomindah Twist, 1962 1962 Maneroo 1952 Roma 1964 Longreach 1955 Jundah

Mines Administration Pty Ltd, 1963 Geological Survey of Q'land, 1960 Mott & Associates, 1964a Geological Survey of Q'land, 1960

W.O.L. No. 3 24°22'45" 142°46'20" (Warbreccan)

1955 Jundah

Geological Survey of Q'land, 1960

N.A.I. Whyen- 28°36'50" 147°21 55 birra No. 1 Alliance Yon- 25°13'19" 143°55'48" gala No. 1

1966 Dirranbandi 1965 Windorah

Cundill, Meyers & Associates, 1967 Laing, 1966

/

,/

* South Australia. Spec.Publs geol.Soc.Aust, 4: pp. 87-118, Pis 1-2, 1973.

115

D. BURGER APPENDIX B

116

Location, stratigraphic position and BMR serial number of rock samples in the Murospora florida Zone Surat Basin BMR Notes Formation/ Zonal affinity No. Depth of Sample Well/Bore Member feet metres (MFP) core Minmi Foraminisporis asymmetricus 56-5 4326 185'5" 64-2 4328 210'7" >> 73-6 4329 241'6" Nullawurt 851 4330 279 4 ' Foraminisporis wonthaggiensis Kingull 93-6 4457 BMR Roma 1 307' 94-8 4458 311' >> 106-1 4460 348' Cicatricosisporites australiensis 1150 4462 Mooga 123-9 4463 406'5" » core Minmi Foraminisporis asymmetricus 99'6" 30-3 4286 Nullawurt 48-0 4288 1578" 'ir 64-9 4289 BMR Foraminisporis wonthaggiensis Kingull 85-8 4278 Mitchell 11 28 r 6" Cicatricosisporites australiensis Mooga 347' 105-8 4279 core Murosporaflorida? Nullawurt/ 38-4 3967 126' Minmi correlate BMR 86-6 3969 Mooga correlate Cicatricosisporites australiensis 284' Mitchell 1 347 5 ' 105-9 3970 core Doncaster ? palynological unit K lb-c? 102'2" 311 4676 Murosporaflorida? Bungil 159'10" 48-7 5131 BMR Roma 8 212'9" 64-8 5132 >> ? 264'2" 80-5 5133 Cicatricosisporites australiensis core Orallo 254-5 880 A.A.O. 1 835' (Roma) core 1582' 482-2 901 Minmi correlate Foraminisporis asymmetricus U.K.A. swc 1848' Cabawin 1 563-3 1177 Kingull correlate Cicatricosisporites australiensis core 2100' 640-1 902 Orallo correlate

„ „ 99

/

/

99

99

2 1 2

99

99

99

99

99

99

99

99

99

99

99 99

/

/

99

99

J>

99

„ 99

99

99

Eromanga Basin Depth of isample BMR No. feet metres (MFP) 55-0 4311 BMR 180'6" 79-6 4313 Mitchell 7 2613" 20-6 4383 BMR 67'9" 65-3 4385 Augathella 3 214'3" 23-7 4181 BMR Tambo 77'ir 5 26-8 4176 srio" 4182 29-0 4183 95' BMR 153' 46-6 3358 Longreach 3 26-2 5545 BMR 86' Jericho 1 62 4335-6 BMR Hughen- 202-5" den 1 Well/bore

Formation/ Member

Zonal affinity

Notes core

99

Foraminisporis wonthaggiensis Cicatricosisporites australiensis Foraminisporis asymmetricus Murospora florida Foraminisporis wonthaggiensis Cicatricosisporites australiensis

Ronlow

Foraminisporis wonthaggiensis

core

Ronlow

Foraminisporis wonthaggiensis

core

Gilbert River

Foraminisporis asymmetricus

core

Hooray 99

Doncaster Hooray Hooray

99

core 99

core 99 99

Well/Bore BMR Gilberton2 F.D. Alice River 1 L.O.L. Saltern Ckl L.O.L. Marchmont 1 Exoil Brookwood 1 A.A.O. Penrith 1 Conorada Ooroonoo 1 A.A.P. Mayneside 1 Amerada Newlands 1 W.O.L. 2 (Warbreccan) W.O.L. 3 (Warbreccan) Alliance Yongala 1 Ph.-S. Cothalow 1 Alliance Chandos1 Smart Oil Orient 2 D.F.S. Innamincka 1 N.A.I. Whyenbirra 1

SPORE ZONATION AND HISTORY OF NEOCOMIAN Eromanga Basin—continued Formation/ Depth of Sample BMR No. Zonal affinity Member feet metres (MFP)

117 Notes

84'8" 25-8 113' 34-4 870' 265-2

5481 5482 2369

Doncaster Gilbert River Hooray

Foraminisporis wonthaggiensis

core

1654'

5543

Hooray

Foraminisporis wonthaggiensis

core

Hooray

504-1

Foraminisporis asymmetricus ? 99

core

2247'

684-9

5544

Cicatricosisporites australiensis

core

1992' 2110' 2754' 2962' 2376' 2578' 3728' 3984'

607-2 643-1 839-4 902-9 724-2 785-8 1136-3 1215-3

2252 Doncaster Foraminisporis asymmetricus ? 2253 Hooray correlate Foraminisporis wonthaggiensis 2223 Hooray Cicatricosisporites australiensis 2224 942 Mesozoic undiff. Foraminisporis wonthaggiensis 1171 Murospora florida Hooray Foraminisporis asymmetricus » Murospora florida

core

3818' 4156' 3542' 3040'

1163-7 1266-7 1079-5 926-6

4114 4104 556 560

Hooray ii Hooray >»

4447'

1355-4

3878

Hooray

Foraminisporis wonthaggiensis

core

3376' 3394' 3630' 4682' 4685' 2200' 2400' 2600' 3938'

1029-0 1034-5 1106-4 1427-1 1428-0 670-6 731-5 792-5 1200-3

5555 5559 5556 4147 4148 5562 5557 5558 309

Hooray »

core

Hooray »

Foraminisporis asymmetricus Foraminisporis asymmetricus ? Murospora florida Foraminisporis asymmetricus

core

Hooray »»

Foraminisporis wonthaggiensis

core

99

99

Hooray

1400' 1600' 1800' 2038'

426-7 487-7 548-6 621-2

4268 4269 4270 5093

Foraminisporis asymmetricus (upper limit) Foraminisporis asymmetricus (upper limit) Foraminisporis wonthaggiensis Cicatricosisporites australiensis

99

Hooray >» 99 99

Spec.Publs geol.Soc.Aust, 4: pp. 87-118, Pis 1-2, 1973.

Cicatricosisporites australiensis 99

Cicatricosisporites australiensis Foraminisporis asymmetricus

99

99

99

99

core 99

core 99

See Poumot (in Zolnai, 1965) core swc core 99

99

99

99 99

core swc 99

core 99

D. B U R G E R EXPLANATION OF PLATES PLATE 1

Stratigraphically significant palynomorphs in the Great Artesian Basin, Queensland. All figures approximately X 550. Fig. 1. Couperisporites tabulatus Dettmann, 1963. BMR Mitchell No. 11; 347 feet (105.8 m), Mooga Sandstone. Prep. 4279-2; 333-1156 (CPC 12168). Fig. 2. Cyclosporites hughesi (Cookson & Dettmann) Cookson & Dettmann, 1959a. Alliance Yongala No. 1; 4,447 feet (1,355.4 m) Hooray Sandstone. Prep. 3878-2; 435-1145 (CPC 12169). Fig. 3. Cicatricosisporites australiensis (Cookson) Potonie, 1956. UKA Cabawin No. 1, 1,582 feet (482.2 m). Minmi Member correlate. Prep. 901-1; 290-1086 (CPC 12170). Fig. 4. Foraminisporis asymmetricus (Cookson & Dettmann) Dettmann, 1963. Alliance Chandos No. 1; 4,682 feet 0,427.1 m), Hooray Sandstone. Prep. 4147-2; 433-1164 (CPC 12171). Fig. 5. Cicatricosisporites hughesi Dettmann, 1963. Alliance Yongala No. 1; 4,447 feet (1,355.4 m), Hooray Sandstone. Prep. 3878-1; 356-1044 (CPC 12172). Fig. 6. Cicatricosisporites ludbrooki Dettmann, 1963. Alliance Chandos No. 1; 4,685 feet (1,428.0 m), Hooray Sandstone. Prep. 4148-1; 293-1060 (CPC 12173). Fig. 7. Crybelosporites stylosus Dettmann, 1963. BMR Roma No. 1; 55 feet (16.9 m), Minmi Member. Prep. 4321-1; 431-1181 (CPC 12174). Fig. 8. Reticuloidosporites arcus (Balme) Dettmann, 1963. BMR Roma No. 1; 348 feet (106.1 m), Kingull Member. Prep. 4460-1; 402-1083 (CPC 12175). Fig. 9. Januasporites spinulosus Dettmann, 1963. BMR Mitchell No. 7; 180 feet (55.0 m), Hooray Sandstone. Prep. 4311-2; 399-1185 (CPC 12176). Fig. 10. Januasporites spinulosus Dettmann, 1963. BMR Roma No. 1; 348 feet (106.1 m), Kingull Member. Prep. 4460-1; 423-1111 (CPC 12180). Fig. 11. Foraminisporis wonthaggiensis (Cookson & Dettmann) Dettmann, 1963. Alliance Chandos No. 1; 4,685 feet (1,428.0 m), Hooray Sandstone. Prep. 4,148-1; 280-1148 (CPC 12177). Fig. 12. Nevesisporites vallatus de Jersey & Paten, 1964. Alliance Yongala No. 1; 4,447 feet (1,355.4 m), Hooray Sandstone. Prep. 3878-1; 352-1196 (CPC 12178). Fig. 13. Densoisporites velatus Weyland & Krieger emend. Krasnova, 1961. BMR Roma No. 1; 307 feet (93.6 m), Kingull Member. Prep. 4457-1; 404-1147 (CPC 12179). Fig. 14. Dictyotosporites speciosus Cookson & Dettmann, 1958. Smart Oil Orient No. 2; 2,200 feet (670.6 m), Hooray Sandstone. Prep. 5562-2; 456-1178 (CPC 12181). Fig. 15. Dictyotosporites speciosus Cookson & Dettmann, 1958. BMR Mitchell No. 7; 180 feet (55.0 m), Hooray Sandstone. Prep. 4311-1; 288-1200 (CPC 12182). PLATE 2

Stratigraphically significant palynomorphs in the Great Artesian Basin, Queensland. Fig. 1. Pilosisporites notensis Cookson & Dettmann, 1958, X 380. BMR Mitchell No. 11; 158 feet (48.0 m), Nullawurt Sandstone Member. Prep. 4288-1; 398-1090 (CPC 12183). Fig. 2. Laevigatosporites sp. X 380. Alliance Yongala No. 1; 4,447 feet (1,355.4 m), Hooray Sandstone. Prep. 3878-2; 351-1109 (CPC 12184). Fig. 3. Cooksonites variabilis Pocock, 1962. X 550 BMR Roma No. 1; 279 feet (85.1 m), Nullawurt Sandstone Member. Prep. 4330-1; 355-1033 (CPC 12185). Fig. 4. Murospora florida (Balme) Pocock, 1961. X 380. BMR Mitchell No. 11; 99 feet (30.3 m), Minmi Member. Prep. 4286-1; 440-1172 (CPC 12186). Fig. 5. Trilobosporites purverulentus (Verbitskaja) Dettmann, 1963. X 380. BMR Roma No. 1; 279 feet (85.1 m), Nullawurt Sandstone Member. Prep. 4330-1; 352-1055 (CPC 12187). Fig. 6. "Gen et sp. indet. forma A" Eisenack & Cookson, 1960. X 550. AAO Penrith No. 1; 2,754 feet (839.4 m), Hooray Sandstone. Prep. 2223-1; 307-1138 (CPC 12188). Fig. 7. Muderongia tetracantha (Gocht) Alberti, 1961. X 380. BMR Mitchell No. 11; 99 feet (30.3 m), Minmi Member. Prep. 4286-1; 293-1038 (CPC 12189). Fig. 8. Canningia colliveri Cookson & Eisenack, 1960. X 380. BMR Mitchell No. 11; 99 feet (30.3 m), Minmi Member. Prep. 4286-1; 422-1178 (CPC 12190). Fig. 9. Dingodinium cerviculum Cookson & Eisenack, 1960. X 550. BMR Augathella No. 3; 68 feet (20.6 m), Doncaster Member. Prep. 4383-1; 379-1032 (CPC 12191). Fig. 10. Odontochitina operculata (Wetzel) Deflandre & Cookson, 1955. X 380. BMR Hughenden No. 1; 205 feet (62.4 m), Gilbert River Formation. Prep. 4336-1; 314-1128 (CPC 12192).

D.

BURGER

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE 1

PLATE

2

D.

BURGER

SPORE-PLANT RELATIONSHIPS IN VICTORIAN MESOZOIC CRYPTOGAMS By J. G. DOUGLAS (With 2 Tables and 13 Text-Figures) ABSTRACT

Megafloral and microfloral components of the cryptogam element in the Lower Cretaceous floras of Victoria, Australia, are discussed; and in situ sporomorphs and their parent plants are briefly described. Among these are Coniopteris and Adiantites spp., several new Filicales species, and a new member of the Hepaticae. INTRODUCTION previously been shown (Douglas, 1969), that The Victorian Mesozoic floras are contained the existence of a large bennettitalean element, in a thick sequence of sediments deposited in various pteridosperm groups, and a diverse an east-west trough in the south of the State ginkgoalean assemblage is not revealed by the and extending offshore. The geology and strati- current interpretation of the microflora. graphy of these beds have been discussed in a TABLE I number of reviews and original papers, inclu- Number of cryptogam taxa suggested by (1) microfloral, ding Edwards & Baker (1943), Singleton (2) megafloral remains and (1967), and Douglas (1969). For many years they were regarded as Jurassic in age, following (2) (1) palaeontological work of Seward (1904), MedMegaflora Sporomorphs well (1954) and others, but Kenley (1954), Fungi 5 Cookson & Dettmann (1958a), and Dettmann Algae 1? (1963) showed that most of the section was of 8-10? 8 Early Cretaceous age. Deposition was almost Hepaticae Musci Many (20?) 1 entirely continental and palynological studies Lycopodiales 1 have been of major importance in effecting Isoetales 4 stratigraphical subdivision. Prominent contri- Sphenopsida 30+ 12? butions in this field have been made by Cook- Filicales son & Dettmann (1958a, 1959), Dettmann On the other hand the megafloral remains (1963), Evans (1966), and Dettmann & Playdo not show the diversification of the Lycoford (1969). which is clearly indicated in the microDouglas (1969) provided a recent examina- podiales Many of the apparent contradictions betion of most of the non-cryptogam elements flora. the ranges of the two components are of the megafloras, and in 1971 described the tween products of their differing natures, but other cryptogams. factors must also be taken into account. For example the absence of megascopic lycopods THE CRYPTOGAMS may be explained to some degree by the major Cryptogam taxa are listed in Table I. It development of this order in the older submust be emphasized that relationships of many surface zones, rather than the younger outof the sporomorphs have not been established, crop areas yielding most of the hand specimens and that figures in this category are speculative. (see Table II). A striking feature of the table is the difThere are other areas in which the evidence ference in the composition of the ancient cryp- of the megafloral and microfloral components togam floras presented by the two components. is in accord. For example Table II shows deOnly in the Hepaticae are the numbers of taxa velopment or evolution within the Lower Crein close agreement. Table I also shows that taceous of certain elements. Megafloras and there is little suggestion in the microfloras of microfloras both show an increase in importhe extensive if taxonomically weak sphenopsid tance of the Hepaticae, and a marked reducelement in the older part of the section. It has tion in the role of the lycopods. In the case of —

—

—

— —

Spec.Publs geol.Soc.Aust, 4: pp. 119-126, 1973.

—

J. G. DOUGLAS 120 the Filicales the contention of Dettmann & the floras of eastern Australia and those of Playford (1969) that they become more pro- western North America, U.S.S.R. and New minent in the younger zones is difficult to Zealand in younger beds of late Albian-early assess in the megafloras. Several new species Senonian (Late Cretaceous) age. certainly appear in the youngest megafloral The major point of the these comparisons zone (Zone D:Douglas, 1969) but it is doubt- and speculations is that information based on ful if this represents an overall increase in the megafloral remains is vital for better underfern representation. Many of the collections standing of the ancient floras. Apart from acafrom the oldest outcrop and sub-surface areas demic considerations, such information has in contain a higher percentage of ferns. the past been used principally to improve stratigraphical knowledge, but it has long been recognized that it is in the field of palaeoTABLE II Development of major cryptogam groups through the environment that the advantages of costudy are greatest. Several authors have pointed out Victorian Mesozoic sequence that the plant body, in constant contact with the atmosphere and shallow subsurface, is perMega Sporomorphs Zone or remains Sub-zone haps the best palaeoenvironment indicator available. From Dettmann & Playford (1969) Presented below is a short survey of sporoParadoxa morphs found in situ and fertile organ-vegeta8CJ Striatus tion relationships observed in the cryptogam o Speciosus CU of the Victorian Lower Cretaceous Hughesi 8 component £Q* Stylosus floras. 1. Fungi and Algae. Fungal spores often do Although lack of knowledge of the meganot survive palynological treatment and are floras of several other Lower Cretaceous areas rarely considered in sporomorph assessmakes comparisons difficult, the megafloral ments unless they have some particular sigevidence also supports the suggestions about nificance or are in unusually large numbers. the palaeogeographic range of the floras made They may however be readily isolated by by Dettmann & Playford (1969). Both comless rigorous maceration techniques. Fertile ponents show striking resemblance between bodies associated with fungi were described eastern Australian bryophytic and pteridofrom the Victorian Lower Cretaceous by phytic (and gymnospermous) elements, and Florin (1952). Douglas (1971) described those of South America and India during Early four Lower Cretaceous fungi but these are Cretaceous (Neocomian—late Aptian) times; small hyphal bodies of no pertinence here. and again there is much resemblance between No alga remains have been identified.

^ Fig. 1. Fig. 2.

1

Figs 1-2. Hepaticites discoides Douglas MS. Planet Tullich Bore 1, 1,540-1,551 ft. GSV 61430, X 1-7. Thallus showing rosette arrangements and dark fertile discs on upper central area. GSV 63903, X 1200. Microspore from appressed mass in fertile disc. Note: GSV refers to Geological Survey of Victoria registered specimen.

SPORE-PLANT RELATIONSHIPS IN MESOZOIC CRYPTOGAMS

Fig. 3.

121

Coniopteris cf. C. hymenophylloides (Brongniart) Seward, X ca 0-8. Reconstruction of sporophyte. The rachis, broken to allow illustration, was longer and more slender than indicated. The rhizome is reminiscent of the present day Pteridium.

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2. Bryophyta. Some indication of a moss element is found in both the megafloras and microfloras, but nowhere is it important. On the other hand hepatics or liverworts are often very numerous and diverse. Fertile organs have been found in situ in one specimen of Hepaticites discoides Douglas MS. from a bore core at 1,540 ft (469 m) in Planet Tullich Bore 1 near Casterton, western Victoria. This plant is a small flat thallus, width 1-2 mm, branching and often radially arranged, forming a hollow rosette. In certain areas, particularly those of thallus junction, there are small, flat, circular, fertile bodies 0.3-0.6 mm diameter, often in groups, with a minor ridge a little inside the periphery. Maceration by standard techniques reveals that these fertile discs contain masses of many thousands of appressed sporomorphs surrounded by a tenuous outer tissue, with major surface indentations. The spores (Fig. 2) are trilete, diameter 35-45^, with prominent surface reticulum forming hexagonal-polygonal areas. Although there is difficulty in reconciling these fertile bodies with the antherideal organ of present day hepatics, this relationship of spore masses to the flat fertile discs has been clearly established. Many small Lower Cretaceous spores with surface reticulum of this nature have been assigned to the Lycopodiales, but these are closely similar to some present day Riccia species. The thallus also suggests mode of growth characteristic of this genus. 3. Lycopodiales. Several small isolated cones from the Koonwarra fish beds (Speciosus Assemblage, or Zone 'C') have tentatively (Douglas, 1971) been allocated to the Lycopodiales. A few of the specimens might yield sporomorphs on maceration, but most are preserved only as impressions, and no spore-cone relationships have as yet been established. 4. Isoetales and Sphenopsida. Although at least 4 sphenopsid species are fossilized and are in great profusion in some Victorian localities, no fertile remains have yet been recognized. Two specimens tentatively allocated to the Isoetales are also sterile. 5. Filicales. Medwell (1954) described only 7 Filicales, but Douglas (1971) distinguished over 30 fern taxa. More than half of these are known only in the sterile condition, but

at least 12 are associated with fertile organs. These include: Sphenopteris warragulensis McCoy Coniopteris cf. C. hymenophylloides (Brongniart) Seward Adiantites lindsayoides Seward, and others described in Douglas (1971) as Coniopteris frutiformis n.sp. Coniopteris nanopinnata n.sp. Coniopteris sp. 'a' Aculea bifida n.sp. Alamatus bifarius n.sp. Amanda floribunda n.sp. cf. Stachypteris sp. and Fern-like foliage sp. 'b' All these possess fern-like fertile organs. There are other ferns where fertile organs are perhaps present, but positive identification is impossible. For example small round areas on certain specimens of the form genus Cladophlebis Brongniart may represent fertile organs. On the other hand, of the ferns listed above, little is known of the fertile organs of S. warragulensis, C. frutiformis, C. nanopinnata, C. sp. 'a,' A. bifida and cf. Stachypteris sp. beyond the shape, size and arrangement of the sori. These suggest relationships with certain present day ferns in some cases. Coniopteris

cf. C. hymenophylloides (Brongniart) Seward Figures 3-6

Stratigraphical range: Zones A-D With the exception of the gametophyte thallus most components of this fern have been found. Both fertile and sterile pinnae were discussed by Seward (1904) and the sterile leaf appears to have been bipinnate or almost tripinnate, and the apices of the pinnae characteristically pointed. In the fertile leaf these apices or pinnules bear caps of flattened hemispherical sori, which in turn contain sporangia 250-300/x in diameter, with annulus cells smallmedium, length 60-80/x. There are 50-100 smooth-walled trilete microspores contained in the sporangia, with often up to 50 per cent aborted. The spores appear to be best listed as Cyathidites australis Couper, common, according to Dettmann (1963), in the Upper Mesozoic of southeastern Australia. There are similar spores described and figured by Tralau (1968) derived from Coniopteris hymenophylloides (Brongniart) Seward f r o m Sweden. The rhizome has been found at one locality attached to the frond.

SPORE-PLANT RELATIONSHIPS IN MESOZOIC CRYPTOGAMS

123

I \

Figs 4-6. Fig. 4. Fig. 5. Fig. 6. Figs 7-8. Fig. 7. Fig. 8.

Figs 4-8 Coniopteris cf. C. hymenophylloides (Brongniart) Seward. NMVP 14193, X 1-7. Jumbunna. Fertile pinna with sori at pinnule segment apices. GSV 63521, X ca 150. Cape Paterson. Sporangium. GSV 63525, X ca 650. Cape Paterson. Microspores removed from macerated sporangium. Note: NMVP refers to National Museum of Victoria registered plant fossil specimen. Adiantites lindsayoides Seward. NMVP 14193, X 1-7. Jumbunna. Fertile pinna with sori on pinnule margins. GSV 63923, X ca 650. Kongwak. Microspores removed from macerated sporangium.

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11

^ 13 Figs 9-13 Figs 9-10. Amanda fioribunda Douglas MS. Casterton. Fig. 9. GSV 63603, X ca 1-3. Fertile foliage fragments. Fig. 10. GSV 63614, X ca 8. Fertile foliage showing distribution of sporangia. Fig. 11. Alamatus bifarius Douglas MS. GSV 60154, X 7. Moonlight Head. Pinnule portions with fertile strips indicated by adhering carbonaceous flakes. Figs 12-13. Fern-like foliage sp. 'b.' Hillside, Glenelg River. Fig. 12. GSV 63535, X 0-8. Leaf with minor indentation. Fig. 13. GSV 63531, X 0-8. Leaf with prominent segmentation.

SPORE-PLANT RELATIONSHIPS IN MESOZOIC CRYPTOGAMS

Adiantites lindsayoides Seward Figures 7-8 Stratigraphical range: Zone C Although this leaf was also described by Seward (1904), less is known about this plant. It is much more stratigraphically restricted than C. cf. C. hymenophylloides but fertile leaf fragments and pinnule segments with marginal sori are common at several Gippsland localities. Pinnule form of new specimens allocated (Douglas, 1971) to this species varies from fanshaped with the arc modified by sori, to bluntly hastate with sori around the apex and along much of the margin. Spores are similar to those from C. cf. C. hymenophylloides and although aborted specimens are much rarer, most sporomorphs have suffered much distortion. These spores are also here tentatively allocated to Cyathidites. Alamatus bifarius Douglas MS Figure 11 Stratigraphical range: Zone D This fern is prominent in new collections from Otway Basin localities. The pinnules are long, straight, and narrow, and in fertile leaves sori are in a row on either side of the main vein. Careful maceration has revealed sporangia. Unfortunately preservation is poor, and sporomorphs have not been isolated. Amanda floribunda Douglas MS Figures 9-10 Stratigraphical range: Zone D The segments of this recently discovered leaf are also long and narrow, although generally shorter and less straight than those of A.

125

bifarius discussed above. The fertile areas are borne on modified pinnules on separate spikes, an arrangement not common in the present day Filicales. Sporangia are prominent, about 300/x in diameter and preliminary examination suggests that the microspores are smooth-walled and trilete. Fern-like foliage sp. 'b' Figures 12-13 Stratigraphical range: Zone D Large leaves, often lobate or partly segmented and fan-shaped, have been found in Zone D localities where the presence of large masses of semi-dispersed or undispersed sporangial clusters of Cicatricosisporites australiensis (Cookson) Potonie (1956) sporomorphs is a characteristic of macerated sediment samples. No closer association of these fossils has been established, but many of the leaves of sp. 'b' are very reminiscent of the present day Schizaea elegans (Vahl.) and close relationship is a possibility. The association of the detached fertile organ called 'fertile organ-bearing microsporangia containing Osmundacidites mollis Cookson & Dettmann,' and the microspores identified under this name was discussed and figured in Douglas (1969). The greater part of the vegetative body of this plant is still unknown. There are other categories of sporomorph fertile organ relationships not considered here. These include the common association of hydropteridean megaspores and certain microspores, often so frequently as to strongly suggest derivation from the same plant. Cookson & Dettmann (19586) and Douglas (1971) comment on several of these.

REFERENCES COOKSON, I. C., & DETTMANN, M. E., 1958a: Some trilete spores from Upper Mesozoic deposits in the eastern Australian region. Proc. R. Soc. Vict., 70, pp. 95-128. , , 19586: Cretaceous 'megaspores' and a closely associated microspore from the Australian region. Micropaleontology, 4, pp. 39-49. , , 1959. Microfloras in bore cores from Alberton West, Victoria. Proc. R. Soc. Vict., 71, pp. 31-38. DETTMANN, M. E., 1963: Upper Mesozoic microfloras from southeastern Australia. Proc. R.

Soc. Vict., 77, pp. 1-148. Spec.Publs geol.Soc.Aust., 4: pp. 119-126, 1973.

, & PLAYFORD, G., 1969: Palynology of the Australian Cretaceous: a review; in Campbell, K. S. W. (Ed.), Stratigraphy and Palaeontology, Essays in Honour of Dorothy Hill, pp. 174-210. A.N.U. Press, Canberra. DOUGLAS, J. G., 1969: The Mesozoic floras of Victoria, Parts 1 and 2. Mem. geol. Surv. Vict.,

28. , 1971: The Mesozoic floras of Victoria, Part 3. Rep. geol. Surv. Vict., 1971/45 [unpublished]. EDWARDS,

A.

B.,

& BAKER,

G.,

1943:

arkose in southern Victoria. Proc. Vict.,

55,

pp.

195-228.

Jurassic

R.

Soc.

J. G. DOUGLAS

126

EVANS, P. R., 1966: Mesozoic stratigraphic paly-

nology in Australia. Aust. Oil Gas J., 12, pp.

58-63. FLORIN, R.,

SEWARD, A. C., 1904: On a collection of Jurassic

plants from Victoria. Rec. geol. Surv. Vict., 1,

pp. 155-211.

1952: On two conifers from the SINGLETON, O. P., 1967: 'South Gippsland' and Jurassic of southeastern Australia. Palaeo'Otway Region'; in McAndrew, J., & Marsden, botanist, 1, pp. 171-182. M. A. H. (Eds), Geology Excursions Handbook. ANZAAS Section C, Melbourne. KENLEY, P. R., 1954: The occurrence of Cretaceous sediments in south western Victoria. TRALAU, H., 1968: Botanical investigations into Proc. R. Soc. Vict., 66, pp. 1-16. the fossil flora of Eriksdal in Fyledalin, Scania. II. The Middle Jurassic microflora. MEDWELL, L. M., 1954: A review and revision oi Sver. geol. Unders. Afh., Ser. C, 633 (Ars the flora of the Victorian Lower Jurassic. 62, 4 ) , pp. 1-185. Proc. R. Soc. Vict., 65, pp. 63-111.

J. G. Douglas, Geological Survey of Victoria, State Offices, 107 Russell Street, Melbourne, Victoria 3000.

RIMULATE POLLEN GRAINS FROM THE LOWER MESOZOIC OF QUEENSLAND By N . J. D E

JERSEY

(With 2 Text-Figures and 5 Plates) ABSTRACT

Four pollen species have been studied, on the basis of features observed with the light microscope and the scanning electron microscope. Most of the material examined came from a sedimentary sequence of Late Triassic to Early Jurassic age, about 1000 m in thickness, in southeastern Queensland. At the generic level, Reyre's broad definition of Classopollis (1970) is accepted. Discisporites is differentiated by its laesurate characters and by the absence of a distal pseudopore. At the specific level, three species of Classopollis—C. chateaunovi, C. simplex and C. meyeriana (Klaus) comb. nov.—are recorded. The diagnosis of the latter, and of Discisporites psilatus de Jersey, 1964, are emended, to include descriptions of the outer sculpture, in Stereoscan view. From the stratigraphic distribution and morphology of the four species, evolutionary relationships among them are suggested. Study of the Queensland material supports Reyre's view that there is progressive development of intrastructure and equatorial striations in Classopollis. Other evolutionary trends in the genus are discussed. A possible origin of rimulate pollen grains of the Classopollis type, via Discisporites, from the Early to Middle Triassic Grebespora, is suggested. INTRODUCTION In a recent paper ( d e Jersey, 1971a) miospore assemblages have been described f r o m a sequence of Lower Mesozoic sediments in southeastern Queensland. T h e sediments are largely, if not entirely, of continental origin; they appear to represent a continuous, conformable sequence about 1000 m in thickness. F r o m the evidence of the assemblages recorded, this thick sequence is regarded as Late Triassic (probably Rhaetian) to Late Liassic in age (de Jersey, 1971a, pp. 24, 2 5 ) . T h e upper part of the succession is correlated with Liassic formations in the Surat Basin, to the west, f r o m which miospore assemblages have previously been recorded by de Jersey & Paten ( 1 9 6 4 ) and Reiser & Williams ( 1 9 6 9 ) . A feature of most of the assemblages f r o m this sequence is the presence of rimulate pollen grains, which have been assigned to the genera Classopollis, Gliscopollis and Discisporites. These grains are characterised by a structure, variously described as a rimula (Pflug, 1953; Pocock & Jansonius, 1 9 6 1 ) , ring-tenuitas (Klaus, 1960) or sub-equatorial circular f u r row (Reyre, 1970). T h e y appear in the basal f o r m a t i o n of the succession (the A b e r d a r e Conglomerate) and become m o r e a b u n d a n t at higher levels; specimens of Classopollis domi-

nate the assemblages f r o m the Helidon Sandstone and M a r b u r g F o r m a t i o n . In this earlier investigation ( d e Jersey, 1971a, pp. 17-19), rimulate grains were assigned to f o u r species—Classopollis classoides Pflug emend. Pocock & Jansonius, 1961, Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969, Discisporites psilatus de Jersey, 1964 and Gliscopollis meyeriana (Klaus) Venkatachala, 1966. Some difficulty was experienced in specific assignment of some of the specimens and some grains were observed which were suggestive of a transition between C. classoides and C. simplex. This led to a fairly broad interpretation of the limits of C. classoides, in distribution studies. A f t e r this study was completed, a paper by Reyre ( 1 9 7 0 ) was received, in which the fine outer sculpture of species of Classopollis (observed with the scanning electron microscope) is described. It is clear f r o m Reyre's w o r k that the surface sculpture of these grains constitutes an additional character which, taken in conjunction with features observed with the light microscope, renders knowledge of their morphology m o r e complete and is thus of considerable value in distinguishing species. Accordingly a study of the fine surface sculpture of the f o u r Queensland species has been

Spec.Publs geol.Soc.Aust., 4: pp. 127-140, Pis 1-5, 1973.

N. J. DE JERSEY 128 made, to provide additional data on the mor- he distinguishes on the basis of this primary phology and relationships of these species and diagnostic character, it is clear that there is to compare them with those described by appreciable variation in the intrastructure, Reyre. The material studied was obtained from presence or absence of equatorial thickening samples examined in this recent work on and development of equatorial striations. Thus material from the Moreton Basin (de Jersey, the features by which Gliscopollis (Klaus) 1971a); reference should be made to this Venkatachala, 1966 has been separated from paper for an account of the stratigraphic and Classopollis, namely the absence of equatorial thickening and striations and minor developgeographic background of the investigation. ment of intrastructure, are shown to be variable, even within individual species. AccordPROCESSING OF SAMPLES AND PREingly the former taxon is included in Reyre's PARATION OF SPECIMENS FOR concept of Classopollis. This interpretation is STEREOSCAN STUDY supported of the Queensland material; Specimens selected for study with the scan- investigationby study species, recorded ning electron microscope were obtained from below, shows ofthatthein four of them there is the maceration residues of samples, previously continuous gradation insomedevelopment of the examined. Reference should be made to an intrastructure and of equatorial thickening. earlier paper (de Jersey, 1971a, pp. 3, 4) for A different view of the circumscription of an account of the methods used in laboratory processing of the samples. Suitable specimens Classopollis was adopted by Medus (1970) selected from these maceration residues were who proposed separation of grains assigned prepared for Stereoscan study according to to the genus, on the basis of Reyre's concept, the methods described by Dettmann (1973, into several different genera. Some of the prop. 8). Microslides of all the specimens, in- posed generic units appear to be of specific, cluding those recovered after study with the rather than generic status. Medus (1970, p. scanning electron microscope, are stored in the 215) argues that the presence of grains of the collection of the Palynology Section, Geo- Classopollis type in different genera of male logical Survey of Queensland. In accordance cones constitutes evidence for subdivision of with previous practice, figured specimens are the dispersed grains into generic, rather than located by the stage co-ordinates of a Carl specific groups. However, there is no necessity Zeiss GFL microscope. Maceration residues for the rank of taxa of dispersed pollen to from all samples are also stored in the collec- correspond to that of the parent cones. A case tion of the Palynology Section and the samples which may readily be cited is that of Cycadothemselves are stored in the Core Library of pites, a genus of dispersed pollen which includes Mesozoic and Tertiary representatives the Geological Survey. of Cycadophyta, Ginkgoales and Pteridospermae. The innate conservatism of the SYSTEMATIC DESCRIPTIONS Cycadopites type of pollen does not permit Genus Classopollis Pflug emend. Reyre, 1970 representatives of these diverse groups to be Type species (here selected): Classopollis distinguished. Similar considerations apply to kieseri Reyre, 1970 Classopollis, and although the disparity in rank of the taxa is not so extreme, species of the Remarks: The generic diagnosis, as emended latter by Reyre (1970, pp. 311, 312) is accepted genera.may well be derived from different cone here. On the basis of this definition, the essenThe distinction of Classopollis from Aporina tial features of the genus are the presence of a sub-equatorial circular furrow (the rimula), Naumova, 1937, and Exesipollenites Balme, a trilete mark and a distal circular pseudo- 1957, has been discussed by Reyre (1970, p. pore. The exine is two-layered; in the inner 309). The former genus has equatorial striaportion of the exoexine various types of intra- tions but lacks a distal pseudopore, rimula and structure are developed. Reyre (1970, pp. 304- trilete mark. Exesipollenites has a distal pseudo308) discusses the use of various characters pore, but no trilete mark or rimula. A genus in differentiating species of the genus and con- which has a distinct rimula is Discisporites cludes, from an impressive volume of evidence, Leschik emend, de Jersey, 1964. However it that the outer sculpture of the grain is the can be distinguished by the absence of a distal ultimate diagnostic character necessary for dis- pseudopore and by the nature of the trilete tinguishing the species. Within species which mark, which consists of simple laesurae, in

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND contrast to the triangular area usually observed in Classopollis. Concerning the type species, Reyre (1970, pp. 309, 310) points out the difficulty of relating his precisely defined species to C. classoides, which was regarded as the type species by Pflug (1953) and Pocock & Jansonius (1961). It should be mentioned here, that as Classopollis was described before January 1st, 1958, proposal of a type species was not necessary to validate the genus (in terms of Article 37 of the International Code of Botanical Nomenclature, see Lanjouw et al., 1966). It is evident from Reyre's discussion that C. classoides has been loosely defined and used in different senses; consequently, in terms of Article 69 of the International Code, it should be rejected as it has been 'used in different senses and so has become a longpersistent source of error'. As the original diagnosis made no mention of the outer sculpture, it is not possible to relate species such as C. chateaunovi Reyre or C. kieseri Reyre to the type material. It is proposed that C. kieseri Reyre, 1970 be selected as the type species to replace C. classoides. This is because, as stated by that author (1970, p. 313) 'C. kieseri resembles C. classoides Pflug, 1953 in many characters visible in the illustrations of this species'. Of the species regarded as valid, it appears closest to the original type material. Thus, even though it does not show well developed striations, it is selected as the type species, following Article 7 of the International Code of Botanical Nomenclature. This states (Note 1): T h e nomenclatural type is not necessarily the most typical or representative element of a taxon; it is that element with which the name is permanently associated'. Botanical affinities: Pollen grains of the Classopollis type have been extracted from male cones by Pettitt & Chaloner (1964), Archangelsky & Gamerro (1968), Archangelsky (1968) and Barnard (1968). Archangelsky assigned his specimens to the Cheirolepidiaceae, which he considered (1968, pp. 164-165) to 'offer a uniform assemblage of plants which may be considered as sufficiently characteristic, being different from other fossil families of conifers mainly in (1) pollen grains, and (2) protection of the ovule by a curvature of the scales'. Barnard concluded more cautiously regarding his contribution (1968, p. 175). This paper goes some way to suggest that we have a coherent group of Jurassic conifers with Brachyphyllum or Pagiophyllum foliage and

129

small male cones bearing Classopollis pollen'. Reyre (1970, p. 317) discusses evidence on affinities, from surface sculpture, and concludes Tn spite of the highly evolved character of the exine, the presence of a proximal trilete scar (sometimes vestigial) is a primitive character. Other Conifers such as Araucariales, Taxodiales and Taxales no longer show any trace of this ancestral character. Therefore plants which produced Classopollis may not be assigned to any Recent order of Conifer ales; they corresponded probably to a special and very large fossil taxon having at least the rank of an order.' There appears, then, to be general agreement on the coniferous affinity of Classopollis, but there is less unanimity on the rank of the group concerned, which is variously assigned to either family or order status. Classopollis chateaunovi Reyre, 1970 (Plate 1, Figs 1, 2, 4, 5, 6; Plate 2, Figs 1-7) Diagnosis (Reyre, 1970, p. 313): 'Subequatorial circular furrow present; trilete scar present, length of laesurae 5-12^; pseudopore diameter 5-10^ intrastructure pseudoreticulate; average exinal thickness 1.5^; equatorial thickening 2.5/*, only vague pseudostriations; band width 8p. Sculpture simple, isomorphous, isodiametric, grumous-verrucose; breadth of grumes 0.2-0.3/*. Size range 20-32,* (100 specimens) '. Figured specimen (Polar view, Plate 1, Figs 1, 2, 4, 5 and 6): Slide S.E.M. 1, stage coordinates 16.3, 92.3; diameter 35,*. Figured specimen (Polar view, Plate 2, Figs 1, 2 and 3): Slide S.E.M. 2, stage co-ordinates 19.0, 89.8; diameter 28,*. Figured specimen (Equatorial view, Plate 2, Figs 4, 6 and 7): Slide S.E.M. 1, stage coordinates 16.1, 91.8; dimensions 23,* (equatorial) x 26fx (polar). Locality of above specimens: G.S.Q. Ipswich 3, 39.8 m (Lower Marburg Formation). Remarks'. The three specimens figured are assigned to this species on the basis of Reyre's diagnosis and illustrations. In particular, the outer sculpture is very similar, as will be evident from comparison of the illustrations, for example, Plate 1, Figure 6 of this paper and Text-Figure 2 (drawing No. 4) of Reyre (1970, p. 320) showing the projections in side view. In the specimen compressed in equatorial aspect (Plate 2, Figs 4, 6 and 7) the equatorial elements of the intrastructure, although largely discontinuous (pseudostriations) are somewhat

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N. J. DE JERSEY

more distinct than in Reyre's illustration (1970, Plate 55, Fig. 11). However, the specimen is considered to indicate slightly more advanced development, within the species, of pseudostriations, because of the close similarity of the outer sculpture (regarded as the ultimate specific character by Reyre) to C. chateaunovi (compare Plate 2, Fig. 7 of this paper and Reyre's Plate 55, Fig. 14). Of the other species described by Reyre, C. bussoni appears closest to C. chateaunovi. It can be distinguished from the latter by its coarser surface sculpture, composed of rugoseverrucose elements which tend to be more elongate and more widely separated. In addition, when studied in equatorial view with the light microscope C. bussoni has well-defined, continuous striations, different from the less complete sub-parallel striations (pseudostriations) of C. chateaunovi. C. chateaunovi can be distinguished from C. meyeriana (Klaus) comb, nov., recorded below, by the psilate character of the exine of the latter, which lacks organized sculptural elements. This distinction is based on Stereoscan study, but it is likely that the species can be differentiated by examination with the light microscope, in the case of specimens preserved in equatorial view. In this aspect specimens of C. chateaunovi exhibit pseudostriations (incomplete striations), as illustrated by Reyre (1970, Plate 55, Fig. 11) and in Plate 2, Figure 4 of this paper, while specimens of C. meyeriana, even those with well-developed intrastructure (as in Plate 3, Fig. 7) lack pseudostriations. In polar view specimens of C. meyeriana with massive intrastructure can also be distinguished from C. chateaunovi, on the basis of the pseudoreticulate to punctate intrastructure of the latter. However, in the Queensland material, a significant proportion of the specimens of C. meyeriana, particularly those from the Helidon Sandstone, have welldeveloped punctate to pseudoreticulate intrastructure, and cannot easily be distinguished from C. chateaunovi with the light microscope. For such specimens Stereoscan study of the surface sculpture provides the only reliable means of specific distinction. In addition to the figured specimens recorded above, specimens with pseudostriations (in equatorial view) are recorded as Classopollis sp. cf. C. chateaunovi. A typical example (refigured from de Jersey, 1970a, Plate 6, Fig. 2) is illustrated in Plate 2, Figure 5 of this paper. Although such specimens have only

been studied with the light microscope, they are regarded as probable representatives of C. chateaunovi. From consideration of the distribution of such specimens, observed in the present study and previously recorded (de Jersey, 1971a, Plate 6, Figs 2, 5) the probable range of the species (in the Moreton Basin) extends through the upper Helidon Sandstone and lower Marburg Formation (Fig. 1, this paper). Distribution: Rhaetian and Hettangian of France (Reyre, 1970, pp. 306-308, 313); lower Marburg Formation of Queensland in G.S.Q. Ipswich 3, at 39.8 m. (this paper). Specimens recorded as Classopollis sp. cf. C. chateaunovi, as discussed above, have also been found in the Helidon Sandstone (G.S.Q. Ipswich 9, 186.7 m) and in the Marburg Formation (G.S.Q. Ipswich 3, 39.8 m. and G.S.Q. Ipswich 5, 27.2 m ) . Similar specimens have been recorded from the Evergreen Formation of the Surat Basin by de Jersey & Paten (1964, Plate 7, Fig. 3) and by Reiser & Williams (1969, Plate 6, Fig. 13). Classopollis meyeriana (Klaus, 1960) comb, nov. et emend. (Plate 3, Figs 5-10; Plate 4, Figs 4-6) Selected synonymy 1960 Circulina meyeriana Klaus, p. 165, Plate 36, Figures 57-60. 1964 Corollina meyeriana (Klaus) Venkatachala & Goczan, p. 129, Plate 3, Figures 1-5, 21. 1966 Gliscopollis meyeriana (Klaus) Venkatachala, p. 99. Emended diagnosis: Subequatorial circular furrow present, ratio of diameter to diameter of amb (in polar view) from about 0.73:1 to 0.81:1. Trilete mark present, in the form of a triangular area with straight or slightly concave sides; sides varying in length from half to about equal amb radius. Distal pseudopore distinct to ill-defined. Intrastructure massive, punctate or pseudoreticulate. Average exinal thickness about 1.5/x, no marked equatorial thickening; no striations. Sculpture psilate to slightly roughened; no organized sculptured elements. Size range (equatorial diameter): 33-40/x (Klaus, 1960); 21-41/x (Queensland specimens—based on 25 measured specimens). Figured specimen (Polar view, Plate 3, Fig. 5): Slide 51827, stage co-ordinates 19.7, 100.1;

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND diameter 30/x (Refigured from de Jersey, 1971a, Plate 6, Figs 6, 7). Locality: G.S.Q. Ipswich 1, 117.2 m. (Ripley Road Sandstone). Figured specimen (Polar view, Plate 4, Figs 4, 5 and 6): Slide S.E.M. 3, stage co-ordinates 12.8, 95.6; diameter 40^. Locality: G.S.Q. Ipswich 3, 362.4 m. Figured specimen (Polar view, Plate 1, Fig. 7): Slide S.E.M. 3, stage co-ordinates 15.5, 97.1; diameter 36/x. Locality: G.S.Q. Ipswich 3, 362.4 m. Figured specimen (Equatorial view, Plate 3, Figs 6, 9 and 10): Slide S.E.M. 6, stage coordinates 15.7, 92.9; dimensions 30/A (equatorial) x 26/JL (polar). Locality: Zlambachgraben bei St Agatha, Upper Triassic of Austria (Rhaetian). Figured specimen (Equatorial view, Plate 3, Fig. 7 ) : Slide S.E.M. 6, stage co-ordinates 13.8, 95.2; dimensions 40/z (equatorial) x 36/x (polar). Locality: G.S.Q. Ipswich 5, 176.4 m. Figured specimen (Equatorial view, Plate 3, Fig. 8): Slide S.E.M. 3, stage co-ordinates 16.2, 96.9; dimensions 41^ (equatorial) x 31/A (polar). Locality: G.S.Q. Ipswich 3, 362.4 m. Remarks: In the above diagnosis, the morphologic terms and order of characters used by Reyre (1970) have been followed. An additional character recorded is the diameter of the rimula (the subequatorial circular furrow), expressed as a ratio to the amb diameter (in polar view); this feature is considered to be of value in distinguishing species in some cases. In earlier studies of the Queensland material this species has been distinguished from specimens assigned to Classopollis classoides by its massive or faintly punctate intrastructure, in contrast to the more definite punctate to pseudoreticulate intrastructure of the latter. This distinction proved difficult to apply, because of the continuous gradation in the intrastructure from massive (as in Plate 3, Fig. 5) to markedly punctate to pseudoreticulate (as in Plate 3, Fig. 7 and Plate 4, Fig. 5). Obviously more precise definition of the species can be achieved on the basis of the outer sculpture of the exine, there being a distinct contrast between the psilate exine of C. meyeriana and grumose-verrucose sculpture of C. chateaunovi, recorded above. Accordingly the specific diag-

131

nosis of C. meyeriana has been emended to include the outer sculpture as a major diagnostic feature. Although the original type material has not been available for Stereoscan study, the emendation is considered justified as specimens from the Upper Triassic of Austria, close stratigraphically and geographically to the type material, have been examined and have a psilate outer surface. In this respect they are similar to the Queensland specimens, which they resemble in all other features. Apart from the variation in intrastructure from massive to punctate or pseudoreticulate, mentioned above, there is appreciable variation in the distinctness of the distal pseudopore. This varies from well defined to barely perceptible. Where evident its diameter is approximately half the amb radius, as indicated by Klaus (1960, p. 165). The species is transferred to Classopollis Pflug emend. Reyre, 1970, as a new combination. This assignment was implied, although not formally proposed by Reyre (1970, p. 309). There is general conformity to the broad diagnosis of that author except that the presence of an inner layer has not been detected. However in other species of Classopollis the layers are often closely attached and the two-layered condition is difficult to observe, except in a few specimens with a shrunken inner layer (e.g. de Jersey, 1971a, Plate 6, Fig. 5). C. meyeriana is distinguished from C. chateaunovi and the other species described by Reyre (1970) by its psilate or slightly roughened outer surface, which lacks any organized sculptural elements. As indicated above, it can also be distinguished from that species, in equatorial view, by the absence of pseudostriations. In extreme cases, specimens of C. meyeriana have a somewhat linear arrangement of the punctate to pseudoreticulate intrastructure at the equator (as in Plate 3, Fig. 7) but do not have a band of pseudostriations. The species differs from C. simplex, recorded below, in lacking the pronounced equatorial thickening of that species, which also has a rimula of larger diameter, relative to the overall diameter of the amb. Discisporites psilatus, also recorded below, lacks a distal pseudopore and, proximally, its simple laesurae are readily distinguished from the triangular area of C. meyeriana. Distribution: Karnian, Norian and Rhaetian of Austria (Klaus, 1960); Rhaetian of Hungary (Venkatachala & Goczan, 1964); Middle and

Spec.Publs geol.Soc.Aust., 4: pp. 127-140, Pis 1-5, 1973.

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N. J. DE JERSEY

Upper Keuper and Liassic of East Germany (Schulz, 1966, 1967); Norian and Rhaetian of West Germany and the Netherlands, Rhaetian of England (Geiger & Hopping, 1968). Queensland records: Aberdare Conglomerate, Raceview Formation, Ripley Road Sandstone, Helidon Sandstone (de Jersey, 1964, 19706, 1971a). In the present study specimens of the species (with psilate outer surface) have been observed in the following samples: G.S.Q. Ipswich 1, 58.5 m. (Ripley Road Sandstone); G.S.Q. Ipswich 3, 161.9 m., 222.5 m., 319.1 m., 352.0 m., 362.4 m. (Helidon Sandstone) and G.S.Q. Ipswich 5, 176.4 m. (Marburg Formation). In accordance with the emendation proposed above a significant proportion of specimens recorded as C. classoides, in assemblages from the Helidon Sandstone (de Jersey, 1971a, pp. 42-45), should be assigned to C. meyeriana. Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969 (Plate 3, Figs 1-4) Description: Subequatorial circular furrow present, ratio of diameter to diameter of amb (in polar view) from about 0.84:1 to 0.90:1. Trilete mark present, in the form of a triangular area with straight or slightly concave sides, about 5 to 8/a in length. Distal pseudopore, diameter 5 to 15^. Intrastructure massive to loosely punctate. Average exinal thickness about IJJL, equatorial thickening 1.5-2/x; no striations. Sculpture psilate to slightly roughened; no organized sculptural elements. Size range (equatorial diameter): 20-40/*, (Danze-Corsin & Laveine in Briche et al, 1963); 18-33^ (Queensland specimens—based on 35 measured specimens). Figured specimen (Plate 3, Figs 1, 2 and 3 ) : Slide S.E.M. 2, stage co-ordinates 18.5, 92.4; diameter 21 p. Locality: G.S.Q. Ipswich 3, 39.8 m. (lower Marburg Formation). Figured specimen (Plate 3, Fig. 4 ) : Slide O.D. 2116, stage co-ordinates 12.9, 90.1; diameter 25/a (refigured f r o m de Jersey & Paten, 1964, Plate 7, Fig. 4 ) . Locality: P.S.Q. Waggaba No. 1 well, sidewall core 9, 1124.1 m. (Precipice Sandstone of Surat Basin). Remarks: Classopollis simplex differs from C. chateaunovi and the other species described by

Reyre (1970) in lacking organized sculptural elements. In this respect it resembles C. meyeriana, described above, but can be distinguished from that species by its equatorial thickening and by the rimula being appreciably closer to the equator, as shown by comparison of the ratios of the diameter of the rimula to the overall diameter of the amb. If further study reveals the presence of intermediate forms and thus demonstrates a gradation between them in these features, there would be justification for regarding C. simplex as a junior synonym of C. meyeriana. The species Classopollis simplex Reyre, 1970 lacks the equatorial thickening of C. simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969 and also differs f r o m that species in having distinct sculptural elements. It is regarded here as a junior homonym of the latter taxon. Although Reyre (1970, p. 308) regards specific diagnoses, which lack a description of the outer sculpture observed by the electron microscope, as invalid, this is not considered by the writer to be applicable to psilate species. Such species must still be differentiated by features observed with the light microscope. The taxonomy of C. simplex has been discussed by Reiser & Williams (1969, p. 16); this interpretation is supported here. These authors also mention the lack of lateral compressions of the species in their material, pointing out that the presence of a thickened equatorial girdle would favour compression in polar orientation. As mentioned recently (de Jersey, 1971a, p. 18) the species Gliscopollis tersus (Norris) Volkheimer, 1968 may be a junior synonym of C. simplex, the only apparent difference being the lack of any record of an intexine in G. tersus. Distribution: Jurassic of France (Briche, Danze-Corsin & Laveine, 1963; Levet-Carette, 1964a, 19646). Queensland records: Precipice Sandstone, Evergreen Formation, Hutton Sandstone of Surat Basin (de Jersey & Paten, 1964; Hill, Playford & Woods, 1966; Reiser & Williams, 1969); Razorback Beds (Playford & Cornelius, 1967); Helidon Sandstone, Marburg Formation (de Jersey, 1971a, this paper). Genus Discisporites

Leschik emend, de Jersey, 1964 Type species: Discisporites niger Leschik, 1955. Remarks: Because Stereoscan study of the Queensland species, Discisporites psilatus, has

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND 13 3 demonstrated its psilate character, it is sus- Locality of above specimens: N.S. 272, 183.9 pected that the type species, D. niger, is also m (Ripley Road Sandstone). psilate and that the granulate appearance described by Leschik (1955, p. 26) is related to Remarks'. Emendation of the specific diagnosis the intrastructure, not the surface sculpture. is proposed, to incorporate a description of surface, observed in Stereoscan study. If this were to be demonstrated by study of the outer is psilate (Plate 4, Fig. 7; Plate 5, Fig. specimens of D. niger with the scanning elec- This 6). There is considerable variation in the width tron microscope, there would be justification distinctness of the rimula. It ranges from for restricting Discisporites to psilate species. and 1.5fi (Plate 4, Fig. 7) down to less than 0.1 Until such Stereoscan study of D. niger is (Plate 5, Fig. 6). In some specimens (e.g., carried out, the generic diagnosis as emended Plate 5, Fig. is accompanied by adjacent by de Jersey (1964, p. 12) is followed. The folds. Another3) itfeature is subject to genus is distinguished from Classopollis Pflug appreciable variation iswhich the intrastructure. emend. Reyre, 1970 by the absence of a distal Specimens observed range from massive (Plate pseudopore and by the character of the trilete 1) to punctate (Plate 5, Figs 2 and 3) mark, which takes the form of long, simple 4,to Fig. pseudoreticulate (Plate 5, Fig. 4). laesurae, in contrast to the triangular area of This species shows a general similarity to Classopollis. the type species, Discisporites niger Leschik, 1955, particularly if, as suggested above, the Discisporites psilatus de Jersey, 1964 emend. 'Komelungskulptur' referred to by Leschik (Plate 4, Figs 1, 3, 7; Plate 5, Figs 1-6) (1955, p. 26) actually represents intrastrucEmended diagnosis: Amb convexly subtrian- ture, not surface sculpture. A distinction can gular to subcircular. Subequatorial circular fur- be made on the basis of overall size, the row present, of variable width, ratio of dia- measurement given by Leschik (23^) being meter to diameter of amb (in polar view) from appreciably below the equatorial diameter of about 0.65:1 to 0.80:1. Trilete mark present, most specimens of D. psilatus. In addition the long simple laesurae, from 2/3 to almost equal rimula appears to be relatively farther from amb radius. No distal pseudopore. Intrastruc- the equator in D. niger, the ratio of its diature massive, punctate or pseudoreticulate. meter to the diameter of the amb being 0.58:1 Average exinal thickness about 1.5/x; no equa- (by measurement from Leschik's figure) as torial thickening; no striations. Sculpture compared with the range of 0.65:1 to 0.80:1 in D. psilatus. Further study of the European psilate; no organized sculptural elements. including an examination of variation Size range (equatorial diameter): 25-41^ species, in these features and of its outer sculpture, (based on 15 measured specimens). is necessary before its relationship to D. psilaFigured specimen (Plate 4, Fig. 1): Slide tus can be accurately assessed. S2410, stage co-ordinates 20.4, 93.4; diameter addition to the specimens recorded above 36jm (Refigured from de Jersey, 1971a, Plate a Inspecimen recently recorded as cf. Disci6, Fig. 10). sporites psilatus (de Jersey, 19716, p. 15, Plate Figured specimen (Plate 4, Figs 3, 7): Slide 4, Fig. 2) is refigured in Plate 4, Figure 2 of S.E.M. 4, stage co-ordinates 15.9, 92.5; dia- this paper. This specimen was found in an meter 38/*. assemblage from N.S. 566, 182.19 m., in the Formation of the Ipswich Coal MeaFigured specimen (Plate 5, Figs 1 and 2): Tivoli sures. It resembles specimens of D. psilatus Slide S.E.M. 4, stage co-ordinates 14.0, 94.1; in having a convexly subtriangular amb, a diameter 40/x. trilete mark with long, simple laesurae and Figured specimen (Plate 5, Fig. 3): Slide punctate intrastructure. However it differs in S.E.M. 4, stage co-ordinates 6.0, 93.5; diameter having a concentric fold instead of a rimula 38/x,. and it is suggested that it represents a tranFigured specimen (Plate 5, Fig. 4): Slide sitional stage between Grebespora concentrica S.E.M. 4, stage co-ordinates 13.9, 93.8; dia- Jansonius and D. psilatus. meter 40ju,. Distribution: Raceview Formation, Ripley Figured specimen (Plate 5, Figs 5, 6): Slide Road Sandstone, Helidon Sandstone of southS.E.M. 4, stage co-ordinates 15.9, 93.5; dia- eastern Queensland (de Jersey, 1964, 19706, meter 36^. 1971a, this paper). M

Spec.Publs geolJSoc.Aust, 4: pp. 127-140, Pis 1-5, 1973.

N. J. DE JERSEY land succession. Thus in Europe, Classopollis meyeriana also precedes C. chateaunovi and C. simplex, as it has been recorded from the Karnian onwards, while the other two species first appear in Rhaetian sediments. Accordingly it is suggested that the two latter species were derived from C. meyeriana, by development of a thickened equatorial girdle in the case of C. simplex and by development of equatorial striations and of surface sculptural elements in the case of C. chateaunovi. The Queensland sequence also provides evidence that this series can be extended to include earlier forms of simpler morphology. A notable example is Discisporites psilatus, which has much in common with C. meyeriana and, like that species, is first recorded from the Aberdare Conglomerate. It is suggested that the latter species arose from D. psilatus by development of a distal pseudopore and of a proximal triangular area, in place of its simple laesurae. Still earlier, a specimen from the Ipswich Coal Measures, recorded as cf. Discisporites psilatus, resembles D. psilatus in all features, except that a distal concentric fold is present in the same relative position as the rimula of that species. In view of the variation in width and distinctness of the rimula in D. psilatus and its association with adjacent folds, such a form seems a likely precursor of that species. Finally, the distinction between the earlier Grebespora concentrica and the Ipswich form lies mainly in the presence of intrastructure in the latter. Consequently a relationship between them is inferred, although MORPHOLOGIC AND STRATIGRAPHIC the apparent absence, in Queensland, of sediRELATIONSHIPS OF THE SPECIES The stratigraphic distribution of the species, ments intermediate in age between the Moolayrecorded above, is indicated in Figure 1. The ember Formation and Ipswich Coal Measures Triassic formations of the Bowen Basin are reduces the prospects of finding intermediate included in this diagram, in addition to the forms. These suggested evolutionary relationships Triassic-Early Jurassic sequence in southeastern Queensland. Relatively continuous ranges are are shown diagrammatically in Figure 2. Evoshown by continuous lines; broken lines indi- lution within the Classopollis group has also cate the sporadic distribution of Discisporites been discussed recently by Reyre, who reached psilatus in the basal Helidon Sandstone and the the following conclusions (1970, p. 320): distribution of specimens recorded as cf. Dis- 41. Classopollis species should be classed accordcisporites psilatus and Classopollis sp. cf. C. ing to the exine sculpture. This system chateaunovi. seems to be expressive of the botanical enFrom consideration of the morphology of tities in the plant group which produced these species it is evident that they form a Classopollis. From our present knowledge, series of increasing complexity, developed by four groups might be suggested, with rugosethe progressive introduction of various characverrucose, echinulate, mixed or double ters. Moreover, three of them have also been structure. recorded from the Late Triassic-Early Jurassic sequence of Europe, where their stratigraphic 2. The palynological evolution inside each group might be different, but it seems that relationship is similar to that in this Queens-

134

Genus Grebespora Jansonius, 1962 Type species: Grebespora concentrica Jansonius, 1962 Grebespora concentrica Jansonius, 1962 (Plate 1, Fig. 3) Diagnosis (Jansonius, 1962, p. 83): 'Size 20 (40 x 45) 55/x, outline circular to subcircular; exine very thin, single-layered, pale, scabrate to laevigate; alete with occasionally a faint suggestion of a rather full Y mark; characteristically with a dark concentric (secondary?) fold near equatorial outline, 1(2) 4^ wide.' Figured specimen: Slide SI667, stage co-ordinates 10.2, 101.4; diameter 26^ (refigured from de Jersey, 1970a, Plate 9, Fig. 9). Locality: D.R.D. 15, 83.3 m (Rewan Formation). Remarks: Jansonius did not indicate the location of the concentric fold in his diagnosis, but from his figure of the holotype (1962, Plate 16, Fig. 3) it is believed to be distal. In Queensland, specimens from the Triassic of the Bowen Basin have been identified with this species (de Jersey & Hamilton, 1967, p. 20; de Jersey, 1968, p. 20; de Jersey, 1970a, p. 19). The specimen figured (Plate 1, Fig 3), which exhibits a vestigial trilete mark, is from the upper Rewan Formation. Although this species is without a rimula, it is included in this paper because of its suggested relationship with Discisporites psilatus, discussed below.

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND 135 the massive infrastructure has preceded into more and more clear and distinct striadifferentiated intrastructure (alveolate, tions. Thus a species with distinct striations punctate, reticulate, etc.). This is organized may be considered highly evolved.' AGE

FORMATION

RHAETO-

FORMATION

RANGE OF SPECIES

MARBURG

-LIASSIC

S»i

HELIDON SANDSTONE RIPLEY ROAD S'STONE

LATE

RACEVIEW

FMN.

Aberdare Conglomerate

TRIASSIC IPSWICH COAL MEASURES

o!

MIDDLE TRIASSIC

MOOLAYEMBER FORMATION CLEMATIS

S'STONE

EARLY

REWAN

TRIASSIC

FORMATION

Fig. 1. Diagram showing ranges of species studied. Age and correlation based on de Jersey (1970a, 1971a, b). Broken lines indicate sporadic distribution, or distribution of comparable forms, as discussed in text. Spec.Publs geol.Soc.Aust, 4: pp. 127-140, Pis 1-5, 1973.

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In terms of the four groups suggested by Reyre, the Queensland species, with psilate or grumous-verrucose sculpture, all fit into his

rugose-verrucose group, which includes 'psilate, grumous, rugose and verrucose sculptures' (Reyre, 1970, p. 318). Within this group, the

C. chateaunovi

RHAET0C. simplex

-LIASSIC

Discisporites

psilatus

LATE Classopollis

meyeriana

TRIASSIC cf. Discisporites

psilatus

MIDDLE

TRIASSIC

EARLY

Grebespora

concentrica

TRIASSIC Fig. 2. Diagram showing suggested evolutionary relationship of species studied. The stippling denotes infrastructure.

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND 137 same pattern of progressive development of land sequence, the first marked development intrastructure and striations is shown in the of a thickened equatorial girdle is in C. simQueensland species. Thus, in the early parts plex; some equatorial thickening is also evident of their ranges specimens of C. meyeriana are in C. chateaunovi. In later species described dominantly massive (Plate 3, Fig. 5). At by Reyre (1970) there is no further develophigher stratigraphic levels, intrastructure of the ment of equatorial thickening. Likewise the punctate and pseudoreticulate types (as in proximal triangular area and distal pseudopore, Plate 4, Fig. 5) is increasingly developed. after their first appearance in C. meyeriana, Eventually, in C. meyeriana, the equatorial do not exhibit progressive modification or elements of the intrastructure become slightly development in later species. Outer sculpture, elongate and attain a sub-parallel arrangement in the form of organized sculptural elements, (Plate 3, Fig. 7) although they cannot be presumably developed from the psilate conclassed as striations or pseudostriations. From dition, which appeared earlier in both Eurosuch forms it is a short step to specimens re- pean and Queensland sequences. Later Jurassic corded as C. chateaunovi and C. cf. chateau- and Cretaceous species described by Reyre exnovi, in which pseudostriations (incomplete hibit diversification of the outer sculpture, relastriations) are developed. These are illustrated tively large spinose projections being developed (in equatorial view) in Plate 2, Figures 4 in some of them. Reyre (1970, p. 318) sugand 5 of this paper. In these specimens many gested that the regional distribution of sculpof the striations are discontinuous; they are tural groups was related to environmental insub-parallel and of varying width. In this re- fluences. Detailed investigation of the surface spect they show a general similarity to the sculpture of Queensland material from higher pseudostriations in Reyre's illustration (1970, Jurassic horizons and from Lower Cretaceous Plate 55, Fig. 11) of C. chateaunovi, pre- sediments will be necessary to determine sumably representing a slightly more advanced whether this hypothesis is also applicable in stage of development of pseudostriations, the Australian region. within the species, than shown by this figure. The remaining aspect of evolutionary deThe final stage in this series consists of the velopment considered in the present study is development of well-defined, continuous, the origin of the rimula itself. As indicated parallel striations. Classopollis grains with stria- above, the suggestion that a simple rimulate tions of this type have recently been observed species, such as Discisporites psilatus, could by Mr J. L. McKellar (pers. comm.) in as- have developed from the Early to Middle Trisemblages from the Hutton Sandstone, in the assic Grebespora concentrica, is supported by Surat Basin. The section containing these the presence in the Ipswich Coal Measures of assemblages is equivalent to the upper part of an intermediate form, recorded as cf. Discisthe Marburg Formation and thus slightly porites psilatus. Further support to this sughigher, stratigraphically, than the top of the gestion is provided by the morphology of section studied in southeastern Queensland (de Paracirculina maliawkinae, described by Klaus Jersey, 1971a). Consequently the entire (1960, pp. 163, 164) from the Karnian of sequence of development of intrastructure and Austria. In this species a rimula and concenstriations can be observed in this Lower Meso- tric fold are both present and closely associated zoic sequence in Queensland; the evolutionary —a relationship favouring the view that the pattern recorded is similar to that described rimula developed as a belt of exinal thinning by Reyre on the basis of material from the adjacent to, and associated with, a concentric Sahara, Israel and France. fold. Other characters which appear to show In conclusion, the present study has proevolutionary development, although less vided some evidence in favour of Reyre's broad clearly and progressively than the intrastruc- generic concept of Classopollis. From the speciture and striations, include the equatorial fic aspect, the occurrence of three European thickening, proximal triangular area, distal species in the Queensland succession extends pseudopore and outer sculpture. Equatorial their known geographic distribution. In addithickening is absent in Discisporites psilatus tion, Reyre's conclusions on the evolutionary and in the early, massive specimens of Classo- development of intrastructure and striations pollis meyeriana. (Some of the later, intra- are supported by data from the Queensland structured specimens of the latter may have material. The local material has also given an slight equatorial thickening). In the Queens- indication as to how rimulate grains of the Spec.Publs geol.Soc.Aust, 4: pp. 127-140, Pis 1-5, 1973.

N. J. DE JERSEY Classopollis type may have developed from versity of Queensland. The writer is indebted simpler forms. It is likely that, in the future, to the officer in charge of the Unit, Mr J. further elucidation of evolutionary trends will Hardy, for assistance during the project. He assist both morphologic and biostratigraphic also gratefully acknowledges the help of his studies in this large and important pollen colleagues in the Palynology Section of the Geological Survey, Dr H. Hekel and Mr J. L. group. McKellar, the former for assistance in preparing specimens for Stereoscan study and the ACKNOWLEDGMENTS latter for assistance in photography and preThe investigation was materially assisted by paration of the text-figures. Mr R. Beaugeais, the advice of Dr Mary E. Dettmann (of the of the Palynology Section, also assisted in Geology Department, University of Queens- photographic work and preparation of the land) on the technique of preparation of pollen plates and text-figures. Professor W. Klaus, of specimens for examination with the scanning the University of Vienna, kindly supplied a electron microscope. This study was carried sample from the Upper Triassic of Austria, to out with the Cambridge Stereoscan instrument, enable Stereoscan study of European specimens of the Electron Microscope Unit, of the Uni- of Classopollis meyeriana. 138

REFERENCES , & HAMILTON, M., 1967: Triassic spores Tomaxeland pollen grains from the Moolayember lia (Coniferae) from the Lower Cretaceous Formation. Pubis geol. Surv. Qd, 336, Palaeof Patagonia (Argentina) and its male and ont. Pap., 10. female cones. J. Linn. Soc. (Bot.), 61 (384), pp. 153-165. , & PATEN, R. J., 1964: Jurassic spores and pollen grains from the Surat Basin. Pubis , & GAMERRO, J. C., 1968: Pollen grains geol. Surv. Qd, 322. found in coniferous cones from the Lower Cretaceous of Patagonia (Argentina). Rev. DETTMANN, M . E., 1973: Angiospermous pollen from Albian to Turonian sediments of eastern Palaeobot. Palynol., 5, pp. 179-182. Australia. Spec. Pubis geol. Soc. Aust., 4, BARNARD, P. D. W., 1968: A new species of Maspp. 3-34. culostrobus Seward producing Classopollis from the Jurassic of Iran. J. Linn. Soc. (Bot.), GEIGER, M . E., & HOPPING, C . A . , 1968: Triassic stratigraphy of the southern North Sea Basin. 61 (384), pp. 167-176. Phil. Trans. R. Soc., Ser. B., 254 (790), pp. BRICHE, P., DANZE-CORSIN, P., & LAVEINE, J. P., 1-36. 1963: Flore infraliasique du Boulonnais HILL, D . , PLAYFORD, G . , & WOODS, J. T., 1 9 6 6 : (Macro-et Microflore). Mem. Soc. Geol. N., Jurassic Fossils of Queensland. Qd Palaeon13. togr. Soc., Brisbane. DE JERSEY, N. J., 1964: Triassic spores and pollen LANJOUW, J., et al. (Eds), 1966: International grains from the Bundamba Group. Pubis geol. Code of Botanical Nomenclature adopted by Surv. Qd, 321. the 10th International Botanical Congress, Edinburgh, August, 1964. International , 1968: Triassic spores and pollen grains Bureau for Plant Taxonomy and Nomenclafrom the Clematis Sandstone. Pubis geol. ture, Utrecht (Regnum Vegetabile, 46). Surv. Qd, 338, Palaeont. Pap., 14. J., 1962: Palynology of Permian and , 1970A: Early Triassic miospores from the JANSONIUS, Triassic sediments, Peace River area, westRewan Formation. Pubis geol. Surv. Qd, 345, ern Canada. Palaeontographica, Abt. B., 110, Palaeont. Pap., 19. pp. 35-98. , 19706: Triassic miospores from the BlackW., 1960: Sporen der Karnischen Stufe der stone Formation, Aberdare Conglomerate and KLAUS, ostalpinen Trias. Jb. Geol. B. A., 5, pp. Raceview Formation. Pubis geol. Surv. Qd, 107-183. 348, Palaeont. Pap., 22. LESCHIK, G . , 1955: Die Keuperflora von Neuewelt , 1971a: Early Jurassic miospores from the bei Basel, II, Die Isosporen und Mikrosporen. Helidon Sandstone. Pubis geol. Surv. Qd, Schweiz. palaeont. Abh., 72, pp. 1-70. 351, Palaeont. Pap., 25. LEVET-CARETTE, J., 1964a: Etude de la microflore infraliassique d'un sondage effectu6 dans le , 19716: Triassic miospores from the Tivoli sous-sol de Boulogne-sur-Mer (Pas-de-Calais). Formation and Kholo Sub-Group. Pubis geol. Annls Soc. geol. N., 83, pp. 101-128. Surv. Qd, 353, Palaeont. Pap., 10.

ARCHANGELSKY, S., 1968: On the genus

RIMULATE POLLEN GRAINS FROM LOWER MESOZOIC OF QUEENSLAND

139

, 1964b: Etude de la microflore bajocienne REISER, R. F., & WILLIAMS, A. J., 1969: Palynod'un sondage effectue dans le sous-sol de logy of the Lower Jurassic sediments of the Roulogne-sur-Mer. Annls Soc. geol. N., 84, northern Surat Basin. Pubis geol. Surv. Qd, pp. 91-121. 339, Palaeont, Pap., 15. MEDUS, J., 1970: Contribution a la classification REYRE, Y., 1970: Stereoscan observations on the des grains de pollen du groupe des Circumpollen genus Classopollis Pflug 1953. Voiles (Pflug) Klaus. Pollen Spores, 12 (2), Palaeontology, 13 ( 2 ) , pp. 303-322. pp. 205-216. SCHULZ, E., 1967: Sporenpalaontologische UnterPETTTTT, J. M., & CHALONER, W. G., 1964: T h e suchungen ratoliassischer Schichten in Zentralultrastructure of the Mesozoic pollen Classoteil des Germanischen Beckens. Paldont. Abh., pollis. Pollen Spores, 6 ( 2 ) , pp. 611-620. Abt. B., 2 ( 3 ) , pp. 547-633. PFLUG, H. D., 1953: Zur Entstehung und Entwick-, in DORING, H., et al., 1966: Erlaiiterungen lung des angiospermiden Pollens in der zur Tabelle der stratigraphischen Verbreitung Erdgeschichte. Palaeontographica, Abt. B., der Sporen und Pollen von oberen Perm bis 95, pp. 60-171. untersten Lias. Abh. Zent. Geol. Inst., 8, pp. PLAYFORD, G., & CORNELIUS, K. D., 1967: Palyno1-203. logical and lithostratigraphic features of the Razorback Beds, Mount Morgan district, VENKATACHALA, B. S., 1966: Mesozoic operculate pollen and their morphology. Palaeobotanist, Queensland. Pap. Dep. Geol. Univ. Qd, 6 (3),

pp. 81-96. POCOCK, S. A. J., & JANSONIUS, J., 1961: T h e

pollen genus Classopollis Pflug, 1953. Micropaleontology, 7 (4), pp. 439-449.

15, pp. 98-101.

, & GOCZAN, F., 1964: The spore-pollen flora of the Hungarian 'Kossen fades'. Acta, geol. Hung., 8 ( 4 ) , pp. 202-228.

N. J. de Jersey, Geological Survey of Queensland, 2 Edward Street, Brisbane, Queensland 4000. EXPLANATION OF PLATES PLATE 1

(All light microscope photographs approximately X 1000 ) Fig. 1. Classopollis chateaunovi Reyre, 1970. S.E.M. polar aspect, distal view showing rimula; pseudopore suggested by faint indentation. X 2500. Fig. 2. Classopollis chateaunovi Reyre, 1970. L.M. (same specimen as Fig. 1) median focus showing rimula, proximal triangular area and punctate-pseudoreticulate intrastructure. Fig. 3. Grebespora concentrica Jansonius, 1962. L.M. polar aspect showing concentric fold and vestigial trilete mark. Fig. 4. Classopollis chateaunovi Reyre, 1970. S.E.M. (same specimen as Fig. 1) showing grumose-verrucose sculpture. X 10,000. Figs 5, 6. Classopollis chateaunovi Reyre, 1970. S.E.M. (same specimen as Fig. 1) showing grumose-verrucose elements in side elevation, adjacent to rimula. X 50,000. Fig. 7. Classopollis meyeriana (Klaus) comb. nov. S.E.M. polar aspect, proximal view showing proximal triangular area. X 2500. PLATE 2

(All light microscope photographs approximately X 1000) Fig. 1. Classopollis chateaunovi Reyre, 1970. S.E.M. polar aspect, distal view showing rimula; pseudopore indicated by distinct indentation. X 2500. Fig. 2. Classopollis chateaunovi Reyre, 1970. S.E.M. (same specimen as Fig. 1) showing grumose-verrucose sculpture (viewed from above). X 50,000. Fig. 3. Classopollis chateaunovi Reyre, 1970. L.M. (same specimen as Fig 1) median focus showing rimula and intrastructure (reduced in area of pseudopore). Fig. 4. Classopollis chateaunovi Reyre, 1970. L.M. equatorial aspect showing punctatepseudoreticulate intrastructure. Adjacent to rimula are sub-parallel striations of variable width, some discontinuous (pseudostriations). (The dark margin is due to heavy gold plating, for S.E.M. study). Fig. 5. Classopollis sp. cf. C. chateaunovi Reyre, 1970. L.M. equatorial aspect. Intrastructure punctate-pseudoreticulate. Imperfect striations, many discontinuous (pseudostriations) adjacent to rimula. Spec.Pubis geol.Soc.Aust., 4: pp. 127-140, Pis 1-5, 1973.

N. J. DE JERSEY Fig. 6. Classopollis chateaunovi Reyre, 1970. S.E.M. (same specimen as Fig. 4). Rimula is indicated by slight indentation; intrastructure and pseudostriations have no surface expression. X 2500. Fig. 7. Classopollis chateaunovi Reyre, 1970. S.E.M. (same specimen as Fig. 4) showing grumose-verrucose sculpture. X 10,000. PLATE 3

(All light microscope photographs approximately X 1000) Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969. S.E.M. oblique polar aspect, proximal view showing equatorial thickening. X 2500. Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969. S.E.M. (same specimen as Fig. 1) showing psilate outer surface. X 20,000. Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969. L.M. (same specimen as Fig. 1) showing rimula (with adjacent folds), equatorial thickening and proximal triangular area. Fig. 4. Classopollis simplex (Danze-Corsin & Laveine) Reiser & Williams, 1969. L.M. polar aspect, median focus showing rimula, equatorial thickenings, proximal triangular area and distal pseudopore. Fig. 5. Classopollis meyeriana (Klaus) comb. nov. L.M. polar aspect, median focus showing rimula, proximal triangular area and distal pseudopore; intrastructure massive. Fig. 6. Classopollis meyeriana (Klaus) comb. nov. L.M. equatorial aspect, showing rimula and distal pseudopore; intrastructure massive. Fig. 7. Classopollis meyeriana (Klaus) comb. nov. L.M. equatorial aspect. Well-developed punctate to pseudoreticulate intrastructure; some of elements are elongated and show sub-parallel arrangement. Fig. 8. Classopollis meyeriana (Klaus) comb. nov. L.M. equatorial aspect, showing rimula and faint punctate intrastructure. Fig. 9. Classopollis meyeriana (Klaus) comb. nov. S.E.M. (same specimen as Fig. 6) showing rimula and distal pseudopore. X 2500. Fig. 10. Classopollis meyeriana (Klaus) comb. nov. S.E.M. (same specimen as Fig. 6) showing psilate (slightly roughened) outer surface. X 20,000.

Fig. 1. Fig. 2. Fig. 3.

PLATE 4

(All light microscope photographs approximately X 1000) Fig. 1. Discisporites psilatus de Jersey, 1964. L.M. polar aspect, median focus, showing rimula and laesurae extending almost to equator. Intrastructure massive. Fig. 2. cf. Discisporites psilatus de Jersey, 1964. L.M. polar aspect, median focus, showing indistinct laesurae, extending almost to equator, and concentric fold. Intrastructure punctate. Fig. 3. Discisporites psilatus de Jersey, 1964. S.E.M. polar aspect; relatively wide, deep rimula. X 2500. Fig. 4. Classopollis meyeriana (Klaus) comb. nov. S.E.M. polar aspect, distal view showing rimula. Pseudopore indicated by slight indentation. X 2500. Fig. 5. Classopollis meyeriana (Klaus) comb. nov. L.M. (same specimen as Fig. 4) median focus showing rimula and punctate-pseudoreticulate intrastructure, absent in area of pseudopore. Fig. 6. Classopollis meyeriana (Klaus) comb. nov. S.E.M. (same specimen as Fig. 4) showing psilate outer surface, adjacent to rimula. X 20,000. Fig. 7. Discisporites psilatus de Jersey, 1964. S.E.M. (same specimen as Fig. 3) showing psilate outer surface and wide rimula. X 20,000. PLATE 5

(All light microscope photographs approximately X 1000) Fig. 1. Discisporites psilatus de Jersey, 1964. S.E.M. polar aspect, distal view showing wide rimula. X 2500. Fig. 2. Discisporites psilatus de Jersey, 1964. L.M. (same specimen as Fig. 1) median focus showing rimula, long laesurae and punctate intrastructure. Fig. 3. Discisporites psilatus de Jersey, 1964. L.M. polar aspect, median focus showing faint punctate intrastructure, fold adjacent to rimula. Fig. 4. Discisporites psilatus de Jersey, 1964. L.M. polar aspect, median focus showing rimula, long laesurae, punctate to pseudoreticulate intrastructure. Fig. 5. Discisporites psilatus de Jersey, 1964. S.E.M. polar aspect, distal view showing narrow rimula. X 2500. Fig. 6. Discisporites psilatus de Jersey, 1964. (same specimen as Fig. 5) showing narrow rimula and psilate outer surface. X 20,000.

N . J . DE JERSEY

Spec.Pubis geol.Soc.Aust., 4, 1973,

PLATE 1

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2

N . J . DE JERSEY

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N . J . DE JERSEY

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J . DE J E R S E Y

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5

REVIEW OF LATE PERMIAN AND TRIASSIC PALYNOLOGY OF NEW SOUTH WALES By ROBIN HELBY (With 4 Text-Figures and 3 Plates) ABSTRACT

Published and unpublished data concerning the distribution of spores, pollen, and microplankton from Late Permian and Triassic strata of New South Wales are reviewed. Comprehensive sequences of Late Permian to Mid-Triassic microfloras are reported from the Sydney Basin and compared with microfloral occurrences below the southern portion of the Surat Basin. Several Early to Mid-Triassic assemblages are recorded from the Lorne Basin. Little useful information is available from the Clarence Basin where carbonisation is normally intense. The abrupt replacement of the Striatites Microflora (Balme, 1964) by the Falcisporites Microflora (formerly Pteruchipollenites Microflora of Balme) is located at the base of the sequence. The subsequent development of the Falcisporites Microflora indicates a strong degree of environmental differentiation with interdigitation of coexistent assemblages. A relatively consistent sequential pattern is evident in the Falcisporites Microflora and five assemblages are recognised. These are in general descending order: 1. The Aratrisporites parvispinosus Assemblage, 2. The Aratrisporites tenuispinosus Assemblage, 3. The Protohaploxypinus samoilovichii Assemblage, ) Comprising a distinct 4. The Lunatisporites pellucidus Assemblage, } Lunatisporites Submicroflora 5. The Protohaploxypinus reticulatus Assemblage. Present data restrict the lower 3 assemblages to the Sydney Basin. The junction of Dulhuntyispora Assemblage Zone (upper division of the Striatites Microflora) and P. reticulatus Assemblage Zone is coincident with the base of the Narrabeen Group, the latter zone occurring in the lower part of the Narrabeen Group. The L. pellucidus Assemblage and the P. samoilovichii Assemblage which comprise the Lunatisporites Submicroflora, exhibit essentially homotaxial although quantitatively distinct microfloras, and occupy the central portion of the Narrabeen Group succession. The A. tenuispinosus Assemblage characterises the upper portion of the Narrabeen Group. A similar microflora is recorded from the Camden Haven Group of the Lorne Basin. The A. parvispinosus Assemblage extends through the Hawkesbury Sandstone and Wianamatta Group in the Sydney Basin and the Wandoan Formation beneath the southern Surat Basin. INTRODUCTION The only published reviews of Australian Late Permian-Triassic microfloral sequences are those of Balme (1964) and Evans (1966). These studies established broadly the framework of the palynological succession, particularly in eastern Australia. Athough palynological data from New South Wales were included in these studies the focus of the authors was directed more specifically to other states. Since compilation of these works extensive drilling both for oil search and for coal exploration has provided an abundance of suitable local material. A number of workers (including Dr N. J. de Jersey, Dr P. R. Evans, and Dr E.

Hodgson) have investigated palynological material related to oil exploration in New South Wales. Their reports are, for the main part, unpublished and I will attempt to summarise and apply these data together with other available information to the expansion and modification of the existing palynological framework. Late Permian and Triassic strata occur extensively in the northeastern quadrant of New South Wales where they are confined within two subparallel north-south trending zones (Fig. 1). The western zone extends along the western edge of the Hunter Thrust system. The sediments in this zone represent the

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DISTRIBUTION OF L A T E P E R M I A N - T R I A S S I C STRATA IN N E W SOUTH W A L E S ( A D A P T E D F R O M 1 : 3 , 0 0 0 , 0 0 0 GEOLOGICAL M A P O F N E W S O U T H

WALES)

153°

Geological Survey of New South Wales Department of Mines Report by R. Helby Date: 20/4/71

149°

5795

Fig. 1.

REVIEW OF LATE PERMIAN AND TRIASSIC PALYNOLOOY OF N.S.W. final fill of the Hunter-Bowen Trough (Newcastle Geosyncline of Voisey, 1959 a, b). A major accumulation of Late PermianTriassic sediments occurs in the central coastal area of the Sydney Basin where the section attains a maximum thickness in excess of 1000 m and ranges in age from Late Permian to Mid-Triassic or slightly younger. In the southern subsurface extension of the Bowen Basin near Moree, 600 m of probable MidTriassic sediments have been intersected in drilling. The eastern zone is composed of the Clarence Basin and the Lome Basin. Although the tectonic relationship of the northeastern areas is uncertain neither appears to include Upper Permian or Lower Triassic strata. Traditionally, the decline and replacement of the Glossopteris flora and its attendant Striatites Microflora (Balme, 1964) are taken as marking the base of the Triassic System in Australia. The coincidence of this horizon with a marked change in the style of sedimentation has reinforced the concept. However, published results of recent microfloral analyses by Evans (1966), Balme (1969) and Helby (1969 ay b; in press) indicate that this event probably occurred within the Late Permian. The coincidence of these major phytostratigraphic and environmental transformations mark this horizon as a logical base for sequential studies of the Triassic. The Triassic-Jurassic boundary in Australia is arbitrarily defined at the first appearance of Classopollis (Banks et al, 1970). In the western zone Triassic sediments are separated from basal Jurassic strata by a substantial paraconformity. The upper limit of Triassic sedimentation is not adequately defined in the Clarence Basin although available data from the Moreton Basin of southern Queensland suggest that the Triassic-Jurassic boundary would be located in the lower part of the Tabulam Group (McElroy, 1962, p. 38). REVIEW OF AVAILABLE DATA It is not possible at this time to integrate fully the palynological data from all areas of New South Wales. For convenience, I will consider information from five principal regions: these are:— 1. The southern and central portion of the Sydney Basin. 2. The northern extension of the Sydney Basin below and adjacent to the southeastern portion of the Great Artesian Basin.

143

3. The southern extension of the Bowen Basin below the Surat Sub-Basin. 4. The Lome Basin. 5. The Clarence Basin. 1. In the Sydney Basin the pioneer work of Dulhunty (1945, 1946) and subsequently Balme & Hennelly (1955; 1956 a, b) established very broadly the nature of microfloral assemblages within the Glossopteris flora. The first microfloral studies beyond the limits of the Glossopteris flora were those of Hennelly. Hennelly (1958a) formally described 7 'microspore' species from a narrow sequence immediately above the Illawarra Coal Measures. His biostratigraphic conclusions were based on 14 productive samples (Hennelly, 1958Z>) and illustrated a substantial and important microfloral change at the base of the Narrabeen Group. He regarded the new species as delineating a Permian/Triassic Transition Zone'. Apart from an investigation of the microfloral sequence across the Newcastle Coal Measures/Narrabeen Group boundary by Grebe (1970), published data from the central and southern portion of the Sydney Basin are syntheses incorporating information from unpublished reports. Grebe's work in the Lake Munmorah area illustrated a considerable microfloral change immediately above the Vales Point Coal Member, which contained a typical Dulhuntyispora Assemblage of Balme (1964). The overlying assemblage contained all the forms described by Hennelly (1958a) and many additional species of spores and acritarchs not known in older microfloras in eastern Australia. The relationship of the Striatiti pollen to the Glossopteris flora in Australia was discussed by Balme (1960). In this presentation he documented the occurrence of Lunatisporites from the Narrabeen Group to the west of Sydney. Similarly, in his outline of Australian pre-Tertiary microfloras, Balme (1964) referred to occurrences of both his 'Taeniaesporites Microflora' and his 'Pteruchipollenites Microflora' in New South Wales strata. In a study which established a palynological framework for Mesozoic sediments in eastern Australia, Evans (1966) delineated 3 major 'palynological units' in the Late Permian/Triassic section. He recognised equivalents of the lower two palynological units and the lower portion of the third unit in the Sydney Basin. The lower palynological unit T r l is divisible in two sub-units, the lower of which is directly comparable to

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Hennelly's Transition Zone. The upper division was characterised by the introduction of Lunatisporites (=Taeniaesporites). Unit Tr2 was dominated by Lunatisporites, the upper subdivision being marked by the appearance of Aratrisporites together with a decrease in the characteristic lycopsid content and a corresponding increase in the abundance of Falcisporites (=Alisporites). Falcisporites dominated unit Tr 3. Evans compared the Early Triassic microflora of the Kockatea Shale, Western Australia (Balme, 1963) with his palynological unit Tr2a, inferring that unit Trl was of Permian age. Helby (1969 a, 6) outlined microfloral successions in the Sydney Basin recognising 4 major assemblages. These syntheses were based on data presented by Helby (1970a) and the relationship of the assemblages to Evans' palynological units can be gauged from Figure 2. Helby suggested that the Permian/Triassic boundary possibly occurred within his Microflora No. 2. Reviewing palynological data in relation to the Permian/Triassic boundary, Balme (1969) concluded that the boundary in Australia was best located at the first appearance of abundant Lunatisporites. Three microfloral assemblages were recognised in an eroded Narrabeen Group sequence to the west of Sydney by Helby (1962). The lower microflora was compared directly to Hennelly's Transition Zone. The overlying microfloras were characterised by abundant Lunatisporites, the middle assemblage being distinguished by the relative abundance of large gymnosperm pollen. The upper assemblage (see Fig. 2) was compared with the microflora of the Kockatea Shale of the Perth Basin, Western Australia and other Late Permian—basal Triassic assemblages; it was concluded that the two lower assemblages of the Narrabeen Group should be referred to the Permian. Although primarily concerned with occurrences in the Bowen Basin, Evans (1963a) indicated an Early Triassic age for the Clifton Sub-group in Strevens Terrigal No. 1 well and suggested, on the basis of microfloral correlation, that the sequence extending from the base of the Gosford 'Formation' to the top of the Wianamatta Group could be compared with the Clematis Sandstone and Moolayember 'Shale' of the Bowen Basin. Helby (1963) referred several assemblages from the A.O.G. Mt Murwin well to the basal Narrabeen Group. A brief outline of the palynostratigraphy of the Narrabeen Group in the Sydney Basin was

presented by Helby (19676). The data were abstracted from accumulating information acquired as part of a systematic study of Sydney Basin microfloras. A more comprehensive treatment of the material is contained in a study by Helby (1970a) which is currently in press. The data from this last study, which were based on 252 productive samples, form the basis of the synthesis discussed below. 2. The northern extension of the Sydney Basin has been comparatively neglected. Helby (1967a) described elements of a microflora from a single sample of the Wollar Sandstone. General aspects of the microfloral sequence in this northern area were also discussed by Hind & Helby (1969). Although palynological data in this area were mainly provided by oil exploration drilling, several outcrop sequences have been examined. Samples from the lower Baltimore Beds in the vicinity of Cobborah yielded typical Mid-Triassic assemblages (Helby, 19656). Three samples extended over 26 m of the Ballimore Beds at Gilgandra (Gilgandra 1:250,000 sheet, G.R. 238 069) yielded microfloras which were referred to Evans' palynological unit Tr3 by Helby (1967c). A pos t-Glossopteris, Late Permian microfloral sequence from the Goulburn River was reported by Helby (1970a). A mid-Triassic assemblage was reported by Hodgson (1963a) from Amoseas Wee Waa No. 1 well. Similar assemblages were recorded in Amoseas Baradine West No. 2 well (Hodgson 19636) and Amoseas Bohena No. 1 well (Evans, 19636). Triassic microfloral assemblages from Alliance Oil Development New Windy No. 1 well (2 samples) and Alliance Petroleum Mirrabooka No. 1 well (4 samples) have been reported by Helby (19666, 1965a respectively). Triassic palynological data from A.O.D. Quirindi No. 1 well and A. J. Woods Poggy No. 1 well are incorporated in Helby (1966c, 1970a). These data, based on nine productive samples, suggest that equivalents of Evans' palynological units Tr2 and Tr3 are represented in these sections. 3. Palynological data from Late PermianTriassic strata of the Bowen Basin in New South Wales are confined to samples obtained from oil exploration drilling below the Surat Basin. All samples occurred within the subsurface Wandoan Formation. De Jersey & Hamilton (1969) published seven microfloral lists of Mid-Triassic forms from the UnionKern-A.O.G. Boomi No. 1 well. A further

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seven assemblages from this well were examined by Evans (1963c), two samples also being recorded by Helby (19716). De Jersey & Hamilton (1969) suggested that the Boomi Wandoan assemblages compared closely with Moolayember Formation microfloras. Similar correlations were suggested by Evans (1963 a, c) and Hind & Helby (1969). Further Wandoan palynological data from U.K.A. Goondiwindi No. 1 well (2 samples: de Jersey, 1964b), U.K.A. Macintyre No. 1 well (2 samples: Helby, 1971a), U.K.A. Esso Gil Gil No. 1 well (18 samples, Helby 1965c, 1966a), and U.K.A. Esso Mt Pleasant No. 1 well (8 samples: Helby, 1966d) have been considered. 4. The microfloras of several outcrop samples from the Camden Haven Group of the Lome Basin were reported by Helby (19706). The assemblages were dominated by Falcisporites spp. but also contained Lunatisporites noviaulensis and Protohaploxypinus samoilovichii together with Aratisporites coryliseminis. A probable upper Early Triassic age was suggested for these assemblages on the basis of comparison with Sydney Basin assemblages. 5. Microfloral data from the ClarenceMoreton Basin are confined to samples from the Nymboida Coal Measures. The quality of the data is further restricted by the relatively high fixed carbon content of these sediments. Brief microfloral lists have been published by de Jersey (1959) and Hennelly (in McElroy 1962, see also McElroy, 1959). I have examined a number of samples from the southern part of the basin, but apart from indicating the presence of the Falcisporites Microflora diagnostic interpretation is prohibited. In following sections references to the successions of microfloras in the Clarence-Moreton Basin are based on the published work of Dr N. de Jersey, who described Late Triassic-Jurassic microfloral sequences of southeastern Queensland. THE MICROFLORAL SEQUENCE Synthesis of the available data reveals a basic sequential pattern in which six and possibly two other consistent microfloral associations are recognised in New South Wales. These associations are designated as assemblages and their vertical and lateral distribution define the relevant assemblage zones. The assemblages are in descending order. 1. The Aratrisporites parvispinosus Assembage, 2. The Aratrisporites tenuispinosus Assemblage,

3. The Protohaploxypinus samoilovichii Assemblage, 4. The Lunatisporites pellucidus Assemblage, 5. The Protohaploxypinus reticulatus Assemblage and, 6. The Dulhuntyispora Assemblage. Dulhuntyispora A S S E M B L A G E The composition of the Dulhuntyispora Assemblage (named by Balme, 1964) has been described by Dulhunty (1945), Balme & Hennelly (1955, 1956a, b), and Evans (1967, 1970). The range of occurrence of the assemblage in New South Wales has been discussed by Dulhunty (1946), Balme (1964), and Evans (1967, 1970). Protohaploxypinus reticulatus ASSEMBLAGE The P. reticulatus Assemblage replaces the Dulhuntyispora Assemblage at a fairly sharp lithological interface immediately above the upper coal measure sequences of the Sydney Basin and its northern extensions. It is not recorded in the southern part of the Bowen Basin in New South Wales, the Lome Basin, or the Clarence Basin. The P. reticulatus assemblage is characterised by the abundance of Falcisporites and by the prominence and occasional abundance of Protohaploxypinus reticulatus (Hennelly). Helby (in press) recorded 73 species of spores and pollen and seventeen species of acritarchs from the P. reticulatus Assemblage Zone. Two zonules can be recognised within the P. reticulatus Assemblage Zone. The lower zonule (Hennelly's Transition Zone) contains a more diverse and usually abundant microflora. Representatives of the Dulhuntyispora Assemblage are still common. Prominent species include Falcisporites australis (de Jersey), Apiculatisporis bulliensis (Hennelly), and Vitreisporites pallidus (Reissinger) Nilsson, 1958. Diagnostic forms include Protohaploxypinus reticulatus, Nevesisporites fossulatus Balme, 1970, Kraeuselisporites ralius Balme, 1970, 'Guthoerlisporites' cancellosus Playford & Dettmann, 1965, Tigrisporites playfordi de Jersey & Hamilton, 1967 and Triquitrites microgranifer Ouyang, 1962. The last five species make their first appearance in this zone. Twenty-two species of spores and pollen, not presently known to occur in the Dulhuntyispora Assemblage, are encountered in this lower P. reticulatus zonule. Acritarchs are also locally abundant: swarms of Quadrisporites horridus

REVIEW OF LATE PERMIAN ANI TRIASSIC PALYNOLOGY OF N.S.W. 147 Hennelly, 1958, Cymatiosphaera Wetzel (Balme) Dettmann, 1963, Lundbladispora emend. Deflandre, 1954 and Micrhystridium brevicula Balme, 1963 and Kraeuselisporites Deflandre emend. Downie & Sarjeant, 1963 cuspidus Balme, 1963. characterising basal Narrabeen Group microOf the 48 species of spores and pollen from floras. this association recorded by Helby {in press), The upper zonule of the P. reticulatus As- 35 species occurred in the underlying P. reticusemblage contains a considerably lower pro- latus Assemblage while 42 species range into portion of Dulhuntyispora Assemblage forms. younger assemblages. Falcisporites dominates these associations while An interpretation of the extent of the L. P. reticulatus and Osmundacidites are promi- pellucidus Assemblage Zone in an approxinent. Lueckisporites nyakapendensis Hart, 1960 mately meridional section through the coastal is confined to this zonule. portion of the Sydney Basin is presented on To the north of Sydney, the P. reticulatus Figure 3. To the south of Sydney this zone Assemblage occurs in the lower, more shaly occurs in the upper portion of the Scarborough portion of the Munmorah Conglomerate; to Sandstone, in the Stanwell Park Claystone, the south of Sydney it occurs in the Coalcliff and in the lower 30 m of the Bulgo Sandstone Sandstone, the Wombarra Shale, and possibly in the N.S.W. Dept Mines Camden D.D.H. 61. the lower portion of the Scarborough Sand- Southwest of Sydney, it occurs in the upperstone; to the west and southwest of the Appin most portion of the Caley Formation and in district it occurs in most of the Caley Forma- most of the lower member of the Grose Sandtion; it has been recognised in outcrop along stone in the N.S.W. Dep. Mines Wollongong the western margin of the Sydney Basin be- D.D.H. 28 and Wollondilly Extended, Northtween Katoomba and Lithgow; to the north ern area D.D.H. 1 and D.D.H. 5. To the north of the Goulburn River this assemblage zone of Sydney it occurs in the upper conglomeratic occurs in at least the basal 20 m of the Wollar facies of the Munmorah Conglomerate and in Sandstone. Its upper limit in this northern area the lower half of the Tuggerah Formation in is not yet defined. Elecom Ourimbah Creek D.D.H. 4, Elecom Ourimbah Creek D.D.H. 5, and in Stevens Lunatisporites SUBMICROFLORA Terrigal D.D.H. 1. The L. pellucidus Assemblage was not enThe L. pellucidus Assemblage and the P. samoilovichii Assemblage can be regarded as countered in samples from north of the Goulcomprising a discrete Lunatisporites (—Tae- burn River, possibly owing to a sample gap niaesporites) Submicroflora within the Falci- due to unsuitable lithologies. Possible palaeosporites Microflora. geographical implications of the absence of this assemblage zone are discussed below. Lunatisporites pellucidus ASSEMBLAGE The L. pellucidus Assemblage is charac- Protohaploxypinus samoilovichii terised by an abundance of L. pellucidus ASSEMBLAGE (Goubin) together with prominent FalciIn contrast to the L. pellucidus Assemblage, sporites and large striate pollen such as Pro- microfloras of the P. samoilovichii Assemblage tohaploxypinus microcorpus (Schaarschmidt) are dominated by pteridophytic forms. At Clarke, 1965, Striomonosaccites morondavensis several localities a transition between the asGoubin, 1965 and Crustaesporites Leschik, semblages is characterised by a sharp decline in 1956. Although the gymnosperm content of the abundance of the large striate gymnosperm the L. pellucidus Assemblage is particularly pollen and corresponding increase in the distinctive the pteridophytic portion of the abundance of a distinctive thick-exined group microflora also displays marked quantitative of pteridophytes together with an increase in changes from the P. reticulatus microflora. the abundance of an undescribed species of Spores characteristic of the Dulhuntyispora Densoisporites. Lunatisporites noviaulensis Assemblage are virtually absent although Dul- (Leschik) and P. samoilovichii (Jansonius) huntyispora parvithola (Balme & Hennelly) Hart, 1964 dominate saccate associations, Potonie, 1956 is relatively persistent in the Falcisporites seldom exceeding two per cent lower range of the zone. Prominent pterido- of that fraction. Aratrisporites makes its first phytic species include Nevesisporites fossulatus, appearance at the base of the assemblage Nevesisporites sp., Retusotriletes radiatus zone and is intermittently dominant in the (Kara Murza), Densoisporites playfordi northern part of the Sydney Basin. Spec.Publs geol.Soc.Aust., 4: pp. 141-155, Pis 1-3, 1973.

ROBIN HELBY 148 Species confined to this assemblage zone 1965, A. tenuispinosus Playford, 1965 and include Rewanisporci foveolata de Jersey, 1970, A. wollariensis Helby, 1967. This zonule has been identified only along Nevesisporites sp. nov., and Welwitschiapites the coastal section of the Sydney Basin and sp. nov. Abundant species include Nevesisporites occupies the sequence between the top of the fossulatus, Retusotriletes radiatus, and Poly- Patonga Claystone and the base of the Bald Hill Claystone. cingulatisporites sp. nov. Sixty-four species of spores and pollen were The upper zonule occupies the portion of the recorded within the P. samoilovichii Assem- Gosford Sub-group extending from the base blage Zone by Helby (in press). Of these at of the Bald Hill Claystone. Falcisporites and least fifty species also occur in older assem- the A. tenuispinosus group are the dominant blages. Of the latter, twenty make their last forms with Lophotriletes novicus Singh, 1964, appearance in this assemblage zone. Nevesisporites limatulus, and Indospora clara South of Sydney the P. samoilovichii As- Bharadwaj, 1962 occurring prominently. The A. tenuispinosus Assemblage was not semblage Zone occupies the major part of the Bulgo Sandstone extending from about 30 m recognised in the northwestern portion of the above the Stanwell Park Claystone to within Sydney Basin. Microfloras from the Camden 15 m or so of the Bald Hill Claystone; west Haven Group of the Lome Basin are broadly of Appin the assemblage occurs in the upper comparable to this assemblage zone athough sandstone member and upper part of the the occurrence of Protohaploxypinus samoilolower sandstone member of the Grose Sand- vichii and Lunatisporites noviaulensis suggest stone; north of Sydney it occupies the upper a possible comparison with the P. samoiloportion of the Tuggerah Formation and the vichii Assemblage or possibly unit Tr2b of Patonga Claystone; to the northwest it is Evans (1966). found in the Digby Beds (equivalent to the lower portion of the Wollar Sandstone), al- Aratrisporites parvispinosus ASSEMBLAGE though the full range of the assemblage has Due to the present absence of suitable yet to be determined in this area. material the precise nature of the change between the A. tenuispinosus Assemblage and Aratrisporites tenuispinosus ASSEMBLAGE the A. parvispinosus Assemblage is not underThe decline of the P. samoilovichii suite stood. The A. parvispinosus Assemblage is and re-emergence of Falcisporites as the particularly diverse and in most samples, dominant saccate component defines the top abundantly represented. Seventy-five species of the P. samoilovichii Assemblage Zone. In of spores and pollen and 10 species of acrithe relatively continuous sequences of the tarchs have been reported (Helby, in press). Sydney Basin this assemblage is replaced by Twenty-eight species not recognised in older the A. tenuispinosus Assemblage. Thirty-nine microfloras occur in this assemblage, 23 of species have been recorded in the A. tenuis- these appearing towards the base of the pinosus Assemblage Zone of which thirty- assemblage zone. The assemblage is dominated three are recognised in older assemblages by Falcisporites and Osmundacidites. I regard (Helby, in press). The dominant elements of the prominent occurrence of Cycadopites the new microflora are Falcisporites, Osmun- follicularis Wilson & Webster, 1946, Protodacidites, and the Aratrisporites tenuispinosus haploxypinus jacobii (Jansonius) Hart, 1964, and Aratrisporites parvispinosus as characgroup. A broader concept of the last-named Two zonules are recognised within the A. teristic. species (Helby, in press; this paper), incorporatenuispinosus Assemblage Zone in the Sydney ting forms having a characteristic exoexine Basin. The lower zonule is particularly dis- structure (with granulate surtinctive. It is dominated by Falcisporites and face expression) predominantly and bearing discrete, scathas very prominent Polypodiisporites muta- tered, mainly echinate includes the bilis Balme, 1970 which is diagnostic of the following species: A. projections, Leschik zonule, Aratrisporites coryliseminis Klaus, emend. Playford, 1965, parvispinosus A. flexibilis Playford 1960, and Aratrisporites tenuispinosus group & Dettmann, 1965, A. paenulatus Playford components. The concept of Aratrisporites Dettmann, 1965 and A. goulburniensis Helby& tenuispinosus applied here is much broader 1967. Other diagnostic species of the assemthan normally accepted. It encompasses forms blage include Verrucosisporites carnarvonensis previously described as A. strigosus Playford,

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REVIEW OF LATE PERMIAN AND TRIASSIC PALYNOLOGY OF N.S.W. 151 de Jersey & Hamilton, 1967, Polypodiaceo- blage Zone and the P. reticulatus Assemblage sisporites sp. nov., Convolutispora sp., Denso- Zone corresponds with a marked sedimentologisporites sp. nov., Duplexisporites problematicus cal change. At this horizon in the Sydney (Couper) Playford & Dettmann, 1965, and Basin twenty-three species of spores and pollen Discisporites verrucosus de Jersey, 1964. make their appearance, and indeed the quantiCadargasporites senectus de Jersey & Hamil- tative and qualitative nature of the microfloral ton, 1967 first appears in higher portions of assemblage change considerably. There is mounting evidence to suggest that the horizon the assemblage zone. The A. parvispinosus Assemblage occurs of this change of microflora represents an in the Hawkesbury Sandstone and Wianamatta erosional hiatus. Shale and their northwestern equivalents in In the northern coastal areas of the Sydney the Sydney Basin. It also occupies the sub- Basin (Fig. 3) there is a complex interdigitasurface Wandoan Formation in the southern tion of assemblages which illustrates the conSurat Basin of New South Wales. However, temporaneous nature of at least three wellthe composition of microfloras from the Wan- delineated, parent floras. I consider that these doan Formation suggest that only higher por- floras were confined to discrete environments tions of the A. parvispinosus Assemblage within the fluviodeltaic system. Their conZone are represented. tinued mutual existence over an extended The Falcisporites Microflora extends be- period is indicated by the occurrence of a yond the upper limit of the A. parvispinosus sample yielding a typical L. pellucidus Assemblage Zone ranging through the Ipswich Assemblage sample immediately above the P. Coal Measures and Bundamba Group in south- samoilovichii Assemblage Zone and a typical east Queensland. However, the relationship P. samoilovichii Assemblage in the A. tenuisof the A. parvispinosus Assemblage to the pinosus Assemblage Zone (Fig. 3). overlying assemblages is not yet known in Balme (1969, 1970) suggested that Early detail. Data presented by de Jersey (1962, Triassic microfloral assemblages with prominent 1964a, 1970, 1971), de Jersey & Hamilton Lunatisporites and the characteristic lycopsid (1965a, b, 1967, 1969) and Evans (1966) microspore suite were restricted and suggest that the assemblages of the Ipswich specialised within a specific coastal environCoal Measures can be readily distinguished ment. The apparent absence of the L. pellufrom microfloras of the overlying Bundamba cidus Assemblage and the increasing promiGroup. Equivalents of these rock units occur nence of Falcisporites in equivalents of the in the Clarence Basin, but their contained P. samoilovichii Assemblage (associations conmicrofloras have not been studied to date. forming to Evans Tr2b unit) to the north of the Goulburn River indicate that the LunaTEMPORAL RELATIONSHIPS OF tisporites Submicroflora lost its identity in SYDNEY BASIN ASSEMBLAGES the 'upland' regions to the northwest of the Central Sydney Basin. Although a general sequence of Late PerThe present, rather sketchy palynological mian and Triassic microfloral assemblages data suggest that the base of the Hawkesbury can be recognised, the detail of the succession is often quite complex (Helby, in press). Sandstone in the Sydney Basin corresponds to Some aspects of this complexity are illustrated the top of the A. tenuispinosus Assemblage on Figure 3, which is an interpretation of Zone. Twenty-seven species of spores and pollen the distribution of the various assemblage first appear in the lower part of the overlying zones. It will be appreciated by comparison A. parvispinosus Assemblage Zone. The rapid of Figures 3 and 4 that the assemblage zones establishment of this number of species suggest a possible depositional hiatus between the are, to varying degrees, facies-dependent. The junction of the Dulhuntyispora Assem- respective zones in the Sydney Basin. REFERENCES Triassic of Western Australia. Palaeontology, bearing on the development of the Glossop6, pp. 12-40. tem-flora; in Leeper, G. W. (Ed.), The Evo—, 1964: The palynological record of Australution of Living Organisms, pp. 269-280. lian pre-Tertiary floras; in Cranwell, L. M. Melbourne Univ. Press, Melbourne. (Ed.), Ancient Pacific Floras, pp. 49-80. , 1963: Plant microfossils from the Lower Univ. Hawaii Press, Honolulu.

BALME, B. E., 1960: Some palynological evidence

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, 1969: The Permian-Triassic boundary in Australia. Spec. Pubis geol. Soc. Aust., 2, pp. 99-112. , 1970: Palynology of Permian and Triassic strata in the Salt Range and Surghar Range, West Pakistan; in Kummel, B., & Teichert, C. (Eds), Stratigraphic Boundary Problems: Permian and Triassic of West Pakistan, pp. 305-453. Kansas Univ. Press, Lawrence. , & HENNELLY, J . P . F . , 1 9 5 5 : Bisaccate sporomorphs from Australian Permian coals. Aust. J. Bot., 3, pp. 89-98. , & HENNELLY, J. P. F., 1956a: Monolete, monocolpate and alete sporomorphs from Australian Permian sediments. Aust. J. Bot., 4, pp. 54-67. , & HENNELLY, J . P . F . , 1 9 5 6 6 : Trilete sporomorphs from Australian Permian sediments. Aust. J. Bot., 4, pp. 240-260. BANKS, M. R. et al., 1970: Correlation charts for the Carboniferous, Permian, Triassic and Jurassic Systems in Australia. I Simposio Internacional sobre Estratigrafia y Paleontologia de Gondwana, Mar del Plata, Argentina, pp. 467-483. DE JERSEY, N. J., 1959: Macro- and micro-floras of north-eastern New South Wales. J. Proc. R. Soc. N.S.W., 92, pp. 83-89. , 1962: Triassic spores and pollen grains from the Ipswich Coalfield. Pubis geol. Surv. Qd, 307. , 1964a: Triassic spores and pollen grains from the Bundamba Group. Pubis geol. Surv. Qd, 321. , 1964b: in Union-Kern-A.O.G. Goondiwindi No. 1 Well Completion Report. UnionKern-A.O.G. [unpublished]. , 1970: Triassic miospores from the Blackstone Formation, Aberdare Conglomerate and Raceview Formation. Pubis geol. Surv. Qd, 348. , 1971: Early Jurassic miospores from the Helidon Sandstone. Pubis geol. Surv. Qd, 351. , & HAMILTON, M., 1965a: Triassic microfloras from the Mount Crosby Formation. Qd Govt Min. J., 66, pp. 324-326. , & HAMILTON, M . , 1 9 6 5 6 : Triassic microfloras of the Moorooka and Tingalpa Formations. Qd Govt Min. /., 66, pp. 327-332. , & HAMILTON, M., 1967: Triassic spores and pollen grains from the Moolayember Formation. Pubis geol. Surv. Qd, 336. , & HAMILTON, M . , 1 9 6 9 : Triassic microfloras from the Wandoan Formation. Rep. geol. Surv. Qd, 31. DULHUNTY, J . A., 1 9 4 5 : Principal microsporetypes in the Permian coals of New South Wales. Proc. Linn. Soc. N.S.W., 70, pp. 147157.

, 1946: Distribution of microspore types in New South Wales Permian coalfields. Proc. Linn. Soc. N.S.W., 71, pp. 239-251. EVANS, P. R., 1963a: Palynological observations on Union-Kern-A.O.G. Cabawin East No. 1 Well, Surat Basin, Queensland. Rec. Bur. Min. Resour. Geol. Geophys. Aust., 1963/21 [unpublished]. , 1963b: (in Gerrard) American Overseas Petroleum Limited, Mid-Eastern Oil N.L. Bohena No. 1, Petroleum Exploration Licence 37, New South Wales, Well Completion Report. American Overseas Petroleum Limited [unpublished]. , 1963c: Lower Mesozoic correlations: Union-Kern-A.O.G. Boomi-Minima-Tingan Wells, Surat Basin. [Unpublished Correlation Chart]. , 1966: Mesozoic stratigraphic palynology in Australia. Aust. Oil Gas J., 12, pp. 58-63. , 1967: Review of the Permian palynology of the Sydney Basin, New South Wales. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1967/103 [unpublished]. , 1970: Upper Carboniferous and Permian palynological stages and their distribution in eastern Australia. I Simposio internacional sobre Estrigrafia y Paleontologia de Gondwana, Mar del Plata, Argentina, pp. 41-54. GREBE, H . , 1970: Permian plant microfossils from the Newcastle Coal Measures/Narrabeen Group boundary, Lake Munmorah, New South Wales. Rec. geol. Surv. NS.W., 12, pp. 125-136. HELBY, R. J., 1962: An introductory study of the palynology of the Narrabeen Group. M.Sc. Thesis, Univ. Sydney [unpublished]. , in

STUNTZ, J . , & WRIGHT, A . J . ,

1963:

Well Completion Report, A.O.G. Mt Murwin No. 1, Sydney Basin, New South Wales [unpublished]. , 1965a: Alliance Oil Development N.L. Mirrabooka No. 1—Interim Report. Palyn. Rep. geol. Surv. N.S.W., 1965/4 [unpublished]. , 19656: Palynological examination of samples from Cobborah. Palyn. Rep. geol. Surv. NS.W., 1965/5 [unpublished]. , 1965c: U.K.A.—Esso Gil Gil No. 1— Interim Report 1. Palyn. Rep. geol. Surv. N.S.W., 1965/3 [unpublished]. , 1966a: U.K.A.—Esso Gil Gil No. 1— Interim Report 2. Palyn. Rep. geol. Surv. N.S.W., 1966/1 [unpublished]. , 19666: Alliance Petroleum Australia N.L. New Windy No. 1. Palyn. Rep. geol. Surv. N.S.W., 1966/3 [unpublished]. , 1966c: Alliance Oil Development Quirindi No. 1—Interim Report. Palyn. Rep. geol. Surv. N.S.W., 1966/9 [unpublished]. , 1966d: U.K.A.—Esso Mt Pleasant No. 1— Interim Report. Palyn. Rep. geol. Surv. N.S.W., 1966/2 [unpublished].

REVIEW OF LATE PERMIAN AND TRIASSIC PALYNOLOGY OF N.S.W. — , 1967a: Triassic plant microfossils from a shale within the Wollar Sandstone, N.S.W. J. Proc. R. Soc. N.S.W., 100, pp. 61-73. — , 19676: A brief review of the stratigraphic palynology of the Narrabeen Group. Palyn. Rep. geol. Surv. N.S.W., 1967/7 [unpublished]. — , 1967c: Palynological examination of outcrop samples from the Gilgandra 1:250,000 sheet. Palyn. Rep. geol. Surv. N.S.W., 1967/1 [unpublished]. — , 1969a: Aspects of stratigraphic palynology in the 'Triassic' of the Sydney Basin. Abstracts of the 1st, 2nd, 3rd, and 4th Symposia on 'Advances in the Study of the Sydney Basin', pp. 17-18. Dep. Geology, Univ. Newcastle. — , 19696: Age of the Narrabeen Group as implied by the microfloras. /. geol. Soc. Aust., 16, pp. 404-405. — , 1970a: A biostratigraphy of the Late Permian and Triassic of the Sydney Basin. Ph.D. Thesis, Univ. Sydney [unpublished]. — , 19706: Plant microfloras from the Lome Basin. Palyn. Rep. geol. Surv. N.S.W., 1970/7 [unpublished]. — , 1971a: Union-Kern-A.O.G. Macintyre No. 1 Well—Interim Report. Palyn. Rep. geol. Surv. N.S.W., 1971/2 [unpublished]. — , 19716: Union-Kern-A.O.G. Boomi No. 1 Well—Interim Reports. Palyn. Rep. geol. Surv. N.S.W., 1971/5 [unpublished]. —, (in press): A biostratigraphy of the Late Permian and Triassic of the Sydney Basin. Mem. geol. Surv. N.S.W.

153

J . P. F., 1958a: Spores and pollen from a Permian-Triassic transition, N.S.W. Proc. Linn. Soc. N.S.W. 83, pp 363-369. , 19586: A palynological investigation of the Transition Zone above the Bulli Seam at Appin No. 4 Bore (Australian Iron and Steel Ltd.). Rep. Coal Res. Sect. CS.l.R.O. Aust., M. 143. HIND, M . C . , & HELBY, R . J., 1 9 6 9 : Geology of the New South Wales portion of the Great Artesian Basin. J. geol. Soc. Aust., 16, pp. HENNELLY,

481-497.

HODGSON, E. A., 1963a: (in Gerrard) American Overseas Petroleum Limited, Mid-Eastern Oil N.L. Wee Waa No. 1, Petroleum Exploration Licence 37, New South Wales, Well Completion Report. American Overseas Petroleum Limited [unpublished]. , 1963 6: (in Gerrard) American Overseas Petroleum Limited, Mid-Eastern Oil N.L., Baradine West No. 2, Petroleum Exploration Licence 37, New South Wales, Well Completion Report. American Overseas Petroleum Limited, [unpublished]. MCELROY, C. T., 1959: Studies in sedimentation and stratigraphy in coal basins in New South Wales—The Clarence Moreton Basin. Ph. D. Thesis, Univ. Sydney [unpublished]. , 1962: The geology of the ClarenceMoreton Basin. Mem. geol. Surv. N.S.W., 17. VOISEY, A. H., 1959a: Australian geosynclines. Aust. J. Sci. 22, pp. 188-198. , 19596: Tectonic evolution of northeastern New South Wales, Australia. / . Proc. R. Soc. N.S.W., 97, pp. 65-72.

R. Helby, 344 Malton Road, Epping, New South Wales 2121.

Spec.Publs geol.Soc.Aust., 4: pp. 141-155, Pis 1-3, 1973.

154

ROBIN HELBY EXPLANATION OF PLATES PLATE 1

(all figures X 500 approx.) Fig. 1. Tigrisporites playfordi de Jersey & Hamilton, 1967 (ranges through the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 2. Cyathidites breviradiatus Helby, 1967 (ranges from the L. pellucidus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 3. Nevesisporites sp. nov. (confined to the P. samoilovichii Assemblage Zone). Fig. 4. 'Nevesisporites' fossulatus Balme, 1970 (ranges from the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 5. Retusotriletes radiatus (Kara-Murza)—(ranges from the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 6. Nevesisporites sp. (ranges from the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 7. 'Nevesisporites' limatulus Playford, 1965 (ranges from the L. pellucidus Assemblage Zone possibly to top of the Falcisporites microflora; see de Jersey, 1971, p. 41). Fig. 8. Apiculatisporis bulliensis (Hennelly) (ranges from the P. reticulatus Assemblage Zone to the P. samoilovichii Assemblage Zone). Fig. 9. Polypodiisporites mutabilis Balme, 1970 (ranges from the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 10. Densoisporites sp. nov. (ranges from the L. pellucidus Assemblage Zone to the A. tenuispinosus Assemblage Zone). Fig. 11. Rewanispora foveolata de Jersey, 1970 (probably confined to the P. samoilovichii Assemblage Zone). Fig. 12. Polycingulatisporites sp. nov. (ranges from the L. pellucidus Assemblage Zone through the P. samoilovichii Assemblage Zone). Fig. 13. Discisporites verrucosus de Jersey, 1964 (ranges from the A. parvispinosus Assemblage Zone into Jurassic assemblages). Fig. 14. Lophotriletes novicus Singh, 1964 (appears in the Middle Permian and ranges through to the A. parvispinosus Assemblage Zone). Figs 15, 16. Triquitrites microgram]er Ouyang, 1962 (probably confined to the P. reticulatus Assemblage Zone). Fig. 17. Polypod iaceosisporites sp. nov. (known only from the A. parvispinosus Assemblage Zone). Fig. 18. Densoisporites sp. nov. (ranges through the A. tenuispinosus Assemblage Zone and the A. parvispinosus Assemblage Zone). Fig. 19. Duplexisporites problematicus (Couper) Playford & Dettmann, 1965 (first appears in the A. parvispinosus Assemblage Zone and ranges into Jurassic strata). Fig. 20. Cadargasporites senectus de Jersey & Hamilton, 1967 (confined to the A. parvispinosus Assemblage Zone). Fig. 21. Densoisporites playfordi (Balme) Dettmann, 1963 (ranges from the P. reticulatus Assemblage Zone to the A. tenuispinosus Assemblage Zone). Fig. 22. lndospora clara Bharadwaj, 1962 (ranges from the Dulhuntyispora Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 23. Cadargasporites sp. nov. (ranges from the P. samoilovichii Assemblage Zone to the A. tenuispinosus Assemblage Zone). Fig. 24. Kraeuselisporites ralius Balme, 1970 (confined to the P. reticulatus Assemblage Zone). Fig. 25. Kraeuselisporites cuspidus Balme, 1963 (ranges from the P. reticulatus Assemblage Zone to the P. samoilovichii Assemblage Zone). Fig. 26. Verrucosisporites carnarvonensis de Jersey & Hamilton, 1967 (confined to the A. parvispinosus Assemblage Zone). Fig. 27. Convolutispora sp. (confined to the A. parvispinosus Assemblage Zone).

REVIEW OF LATE PERMIAN AND TRIASSIC PALYNOLOGY OF N.S.W. PLATE 2

(all figures X 500 approx). Figs 1,2,5. Aratrisporites parvispinosus group (ranges from the A. parvispinosus Assemblage Zone through the Ipswich Coal Measures—de Jersey, 1970). Figs 3.4. Aratrisporites coryliseminis Klaus, 1960 (ranges from the P. samoilovichii Assemblage Zone to the A. parvispinosus Assemblage Zone). Figs 6, 7, 8 Aratrisporites tenuispinosus group (ranges from the P. samoilovichii Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 9. 'Guthoerlisporites' cancellosus Playford & Dettmann, 1965 (ranges from P. reticulars Assemblage Zone into the Bundamba Group—de Jersey, 1970). Fig. 10. Protohaploxypinus reticulatus (Hennelly) ranges from the Dulhuntyispora Assemblage Zone to the A. parvispinosus Assemblage Zone. Fig. 11. Striomonosaccites morondavensis Goubin, 1965 (ranges from the L. pellucidus Assemblage Zone to the P. samoilovichii Assemblage Zone). Fig. 12. Crustaesporites sp. (ranges from the P. reticulatus Assemblage Zone to the P. samoilovichii Assemblage Zone).

PLATE 3

(all figures X 500 approx.) Fig. 1. Protohaploxypinus sp. cf. P. jacobii (Jansonius) Hart, 1964 (confined to the A. parvispinosus Assemblage Zone). Fig. 2. Lunatisporites noviaulensis (Leschik)—(ranges from the L. pellucidus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 3. Lunatisporites pellucidus (Goubin)—(ranges through the L. pellucidus Assemblage Zone to the P. samoilovichii Assemblage Zone). Fig. 4. Protohaploxypinus samoilovichii (Jansonius) Hart, 1964 (ranges from the P. reticulatus Assemblage Zone to the A. parvispinosus Assemblage Zone). Fig. 5. Protohaploxypinus microcorpus (Schaarschmidt) Clarke, 1965 (ranges from the Dulhuntyispora Assemblage Zone to the P. samoilovichii Assemblage Zone). Fig. 6. Lueckisporites nyakapendensis Hart, 1960 (confined to the upper zonule of the P. reticulatus Assemblage Zone). Figs 7, 8, 9. Falcisporites australis (de Jersey)—(extends throughout the Falcisporites microflora). Fig. 10. ? Micrhystridium sp. (confined to the basal P. reticulatus Assemblage Zone). Fig. 11. ? Cymatiosphaera sp. (confined to the basal P. reticulatus Assemblage Zone). Fig. 12. ? Veryhachium sp. (confined to the basal P. reticulatus Assemblage Zone). Fig. 13. Cycadopites follicularis Wilson & Webster, 1946 (first identified in the A. parvispinosus Assemblage and ranges into younger strata). Fig. 14. Welwitschiapites sp. nov. (confined to the P. samoilovichii Assemblage Zone). Fig. 15. Vitreisporites pallidus (Reissinger) Nilsson, 1958 (ranges from Permian to Recent strata). Fig. 16. Quadrisporites horridus Hennelly, 1958 (ranges from the Striatites Microflora to at least the A. parvispinosus Assemblage Zone). Note: Full details of repository and slide locations of the specimens illustrated here is included in Helby {in press).

Spec.Publs geol.Soc.Aust., 4: pp. 141-155, Pis 1-3, 1973.

ROBIN

HELBY

PLATE

25 Spec.Pubis geol.Soc.Aust., 4, 1973.

1

PLATE

2

ROBIN

HELBY

ROBIN HELBY

Spec.Publs geol.Soc.Aust., 4, 1973.

PLATE 3

STUDIES OF NON-CALCAREOUS MICROPLANKTON

TERTIARY NON-MARINE DINOFLAGELLATE CYST ASSEMBLAGES FROM AUSTRALIA By WAYNE K. HARRIS (With 1 Text-Figure and 2 Plates) ABSTRACT

Dinoflagellate cyst assemblages from Tertiary non-marine sediments from Australia are described, illustrated, and compared with those from sediments immediately preceding the Late Eocene transgression in South Australia. The non-marine cyst types are quite distinct morphologically from those in the 'marginal marine' environment. Their diversity is low but they may predominate over other acid-insoluble microfossils. No acritarchs have been observed in any of the non-marine sediments. In contrast, assemblages from 'marginal marine' sediments carry acritarch and dinoflagellate cyst species that are common also to the assemblages recovered from sediments of the open marine environment. Low species diversity and relative frequency are the only factors that may be common to both types of assemblages. Three new genera, Saeptodinium, Morkallacysta and Cubiculosphaera, and five new species, S. gravattensis, S. tasmaniensis, M. pyramidalis, C. maslinensis, and Cleistosphaeridium tenuum, are described. Saeptodinium (al. Peridinium) hansonianum (Traverse, 1955) is a new combination. INTRODUCTION Frequently the palynologist is confronted with the problem of giving a definite answer to the question 'is the sample marine or nonmarine' or 'what is the environment of deposition?' It is relatively simple to provide an answer to the extremes of an intergrading sequence. The open marine sediment will have obvious biological and lithological characters that will distinguish it from its counterpart at the opposite end of the scale, the lacustrine or fluviatile sediment. But it is the fascinating and subtle environmental changes between these limits that provide the palynologist with interpretative difficulties. In this paper I will examine the occurrence and characteristics of some non-marine dinoflagellate cyst assemblages and compare them with assemblages from two transgressive Eocene sequences, one in the Gambier Embayment of the Otway Basin, the other in the St Vincent Basin. Few environmental studies of organic-walled microplankton like those of Wall (1965) on Lower Jurassic microfossils in Britain and of Staplin (1961) on reef-controlled distribution of Devonian microplankton in Alberta, Canada, have been attempted in either marine or non-marine Tertiary sediments. Gocht's recent (1969) monograph on Lower Tertiary assemblages from northwest Germany however

does provide useful data on distribution and frequency in the sections studied. There has been a very strong and understandable bias towards taxonomic and biostratigraphic studies. References to fossil non-marine dinoflagellate assemblages are even fewer and have been almost entirely concerned with Quaternary sediments. Harland & Sarjeant (1970) and Norris & McAndrews (1970) have recently summarised research on this group both fossil and Recent. Important non-marine Quaternary assemblages from Australia have been described by the first two authors and by Churchill & Sarjeant (1962, 1963) and Harland (1971). Harland & Sarjeant (1970) have noted that the only reported previous occurrence of fossil acritarchs in non-marine assemblages has been by Churchill & Sarjeant (op. cit.) and by Sarjeant & Strachan (1968) and these have been in Quaternary sediments. Although the life cycle, and in particular cyst formation, is known for several species of marine dinoflagellates (Evitt & Davidson 1964; Evitt & Wall, 1968; Wall & Dale, 1967', 1968, 1970; Wall, Guillard & Dale, 1967), the non-marine members, with the exception of Peridinium limb at um (Stokes) Lemmermann 1900 from sub-Recent lake muds (Evitt & Wall, 1968) and Ceratium hirundinella (Muller) Schrank (Huber & Nipkow, 1922, 1923; Evitt, 1970), have received little atten-

Spec.Publs geol.Soc.Aust., 4: pp. 159-166, Pis 1-2, 1973.

HARRIS tion despite the fact that they are commonly found today in lakes, reservoirs and even in the laboratory water supply! While less diverse than marine forms they are relatively common; Smith (1950) recorded ten genera from the United States. It is generally agreed that fossil dinoflagellate cysts appear to be most abundant and most diverse in marine sediments and much less abundant or absent in brackish water and non-marine rocks. NON-MARINE TERTIARY ASSEMBLAGES Dinoflagellate cysts have been recovered from several Tertiary sedimentary basins throughout Australia: in Queensland, the City of Brisbane area; in Tasmania, the Mt Bischoff area; in South Australia, the Frome Embayment and the Lake Eyre region of the Great Artesian Basin, and the St Vincent Basin; the central Murray Basin in South Australia and Victoria. City of Brisbane area, Queensland Harris (1965) reported two cyst types from borehole NS5 as Micrhystridium sp. (Cleistosphaeridium tenuum sp. nov.) and aff. Defiandrea sp. (Saeptodinium gravattensis gen. et sp. nov.). The top sample at 130 ft (39.6 m) lies within the Corinda Formation and is a pyritic mudstone. Houston (1965) reported a nonmarine fauna of Pallimnarchus pollens De Vis, 1886 (crocodilian), Ceratodus fosteri Krefft, 1870 (dipnoid), and indeterminate crocodilian, chelonian, fish remains and molluscs. The flora comprises, besides spores and pollen, indeterminate dicotyledonous leaves. The lower sample, 270 ft (82.3 m) is from the Darra Formation and this formation too has a fauna and flora of non-marine aspect that includes bone fragments, an indeterminate mutelid bivalve, seeds, fern pinnules, and dicotyledonous leaves. The age of these formations is Palaeocene (Harris, 1965 and unpublished data). Dinoflagellate cyst frequency varies from 10 to 70 per cent. Great Artesian Basin} South Australia Lower Tertiary sediments are widespread and have been named the Murnpeowie Formation. They consist of fluviatile sands and lacustrine mudstones with abundant leaf impressions. No marine faunas have been reported. The lower unit with dinoflagellate cysts is of Middle to Late Palaeocene age (author's unpublished data). S. gravattensis and MorkaJlacysta pyra-

midalis gen. et sp. nov. comprise less than five per cent of the palynomorph assemblage. Murray Basin, South Australia and Victoria Palaeocene sediments of the Renmark Beds occupy only the central and deepest portion of this basin and have been observed in a number of wells from Waikerie to Loxton and A.A.O. Morkalla No. 1. The lithotopes are fine sands, silts and mudstones and no marine faunas have been reported. Sediments with non-marine dinoflagellate cysts are of Middle to Late Palaeocene age (author's unpublished data). The dinoflagellate assemblage consists primarily of two species, S. gravattensis and M. pyramidalis, the latter being more common. Together their frequency varies from less than one per cent in some samples from Waikerie to more than 45 per cent in Morkalla No. 1. St Vincent Basin, South Australia Sand quarrying at Maslin Bay south of Adelaide exposed a clay lens containing excellently preserved leaves, fruits, flowers, fungi, and mites. The faunal and floral constituents have been listed by Lange (1970), and McGowran, Harris & Lindsay (1970) have determined the age as early Middle Eocene. The lens is interbedded with fluviatile sands and there is no marine fauna. Only one species of dinoflagellate cyst, Cubiculosphaera maslinensis gen. et sp. nov., has been recovered in percentages of the order of five per cent. Mt Bischoff, Tasmania Mudstones beneath basalt at Mt Bischoff have yielded only one species, Saeptodinium tasmaniensis sp. nov., in a frequency of 17 per cent. Well-preserved leaf remains are present but no marine fauna has been reported. The age is Late Oligocene or Early Miocene (author's unpublished data). Summary of characters of the assemblages a. No acritarchs have been observed in any of the samples. b. The cyst types as described here are quite distinct in one or more morphological features such as archeopyle development or wall character. These types have not been encountered in any marine sequence of comparable age in Australia. c. Both capsulate and non-capsulate peridinioid forms are present.

TERTIARY NON-MARINE DINO] d. While species diversity is low, relative frequency with respect to other microfossils is very variable and dinoflagellate cysts may be the dominant fossil group. e. The evidence, both lithological and palaeontological, is not inconsistent with the thesis that they are clearly of nonmarine origin. 'Marginal marine9 assemblages Within South Australia the major Eocene transgression which culminated in the deposition of marl facies of the Lacepede Formation in the Gambier Embayment of the Otway Basin and the Blanche Point Marls of the St Vincent Basin was preceded by clastic

AGELLATE CYST ASSEMBLAGES

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deposition which can be best described as 'marginal marine' in the sense of Taylor (1964). Marginal marine sediments are those deposited 'on the margins between land and sea, such as deltas, estuaries, lagoons and bays' (Taylor, op. cit.). Several bores in the Adelaide city area (loc. 9 on Fig. 1) and in the northern part of the St Vincent Basin at Pt Clinton (loc. 8), where the expression of the transgression is least, yield a characteristic cyst and acritarch assemblage in marginal marine sediments. The stratigraphy and foraminiferal micropalaeontology of the Adelaide area has recently been published by Lindsay (1969). Assemblages are characterised by low species diversity and by a

1000 1 1500 Tasmania \j5l/H06A«T Fig. 1.

Map of localities mentioned in text. 1. Lake Eyre Bore 20 2. E. A. Rudd Bore 5, Frome Embayment 3. Waikerie 27W Bore 4. A.A.O. Morkalla No. 1 Well

5. 6.

7. 8.

9.

Spec.Publs geol.Soc.Aust., 4: pp. 159-166, Pis 1-2, 1973.

Maslin Bay Brisbane City area Mt Bischoff Pt Clinton Adelaide City area

WAYNE I

162

low (five percent) microplankton/spore-pollen ratio. Components include Wetzeliella spp., Defiandrea phosphoritica Eisenack, 1954, Cordosphaeridium sp., Cymatiosphaera sp., and Paralecaniella indentata (Deflandre & Cookson) Cookson & Eisenack, 1970, with cavate cysts dominating. A similar pattern is emerging f r o m the study of sediments of the Lacepede Formation in Observation Bores I & II (Hds of Townsend and Ross respectively, loc. 10) in the Gambier Embayment. Marine influence is marked by the first appearance up section of Defiandrea phosphoritica, Wetzeliella glabra Cookson, 1956, aff. Gonyaulacysta sp., Cymatiosphaera sp., and Paralecaniella indentata. In summary the 'marginal marine' assemblages are characterised by a. low frequency with respect to spores and pollen, b. low diversity, increasing with increasing marine influence, c. presence of acritarchs, d. dominance of cavate and heavily sculptured proximate over chorate cysts, e. the recurrence of constituent species in open marine sediments higher in the successions. Thus non-marine and marginal-marine cyst assemblages would appear to have only one point in common: that is, they may have a low species diversity and a low relative frequency. On all other points the assemblages are distinctive. Non-marine descriptions

microplankton

cyst taxonomy

and

Several of the species described herein have characters which ally them with modern genera such as Peridinium Ehrenberg, 1832, but they lack the characteristic features, such as reflected tabulation, which would enable a firm placement in an extant genus. They are therefore described as cyst form genera and species. The descriptive terminology is that of Downie & Sarjeant (1966). All figured specimens and holotypes are housed in the Palynological Collection of the Geological Survey of South Australia (catalogue numbers prefixed P ) . 'E-W' and 'N-S' coordinates are f r o m Leitz Orthoplan microscope No. 715494 in that institution.

HARRIS Genus Saeptodinium

no v.

Diagnosis: Cavate dinoflagellate cysts of oval to peridinioid form with weakly developed apical and antapical horns. Endocorpus ovoidal. Pericorpus lacking tabulation and with indistinct trace of cingulum. Archeopyle formation not known. Ornament smooth to finely granulate. Type Species: Saeptodinium gravattensis sp. nov. Palaeocene; Queensland. Remarks: This genus closely resembles another freshwater monotypic genus, Teneridinium Krutzsch, 1962, which is much larger and has well developed apical and antapical horns. Krutzsch does not describe an archeopyle and one is not obvious in the photomicrographs. Krutzsch's genus needs further study in the light of more recent studies. Saeptodinium resembles Defiandrea but does not have the characteristic archeopyle of this genus. Content: Saeptodinium hansonianum (Traverse, 1955) f r o m the Brandon Lignite is here transferred to this genus f r o m Peridinium on the basis of its morphology. Saeptodinium

gravattensis

sp. nov.

(Plate 1, Figs 1-4) Diagnosis: A Saeptodinium of ovoid or peridinioid form. Endocorpus ovoidal, enclosed by the pericorpus which has a bulge on the epitract and two slight subequal bulges at the antapex. Periphragm smooth as is the endophragm. The helicoid laevorotary girdle is relatively wide. Archeopyle not observed. Holotype: Plate 1, Figure 2. Darra Formation, Palaeocene. Queensland. Description: Both the periphragm and endophragm are smooth to scabrate and do not show any features of tabulation or any indication of archeopyle formation. The apices of many specimens bear a thickened plug, the function of which is not known. The degree of bulge, and the accompanying indentation in the antapical region is variable f r o m almost no bulge to forms where the indentation is 2-5fi deep. The epitract is subrounded but does show some development of an apical horn. The epitract and hypotract are of approximately equal sizes. The pericoel is 3-5/A wide and wider near the apical region. The girdle is 5-8^ wide and the sulcal groove 5-8/x wide. The periphragm and endophragm are of approximately equal thickness, 0.5//,.

TERTIARY NON-MARINE DINOFLAGELLATE CYST ASSEMBLAGES Dimensions: (10 specimens) Pericorpus: length 50-62/* breadth 45-60/* Endocorpus: length 49-55p. breadth 37-50/*. Remarks: This cyst type is unusual in that no archeopyle has been observed in any specimens. Some specimens have been observed without the apical plug and part of the surrounding wall. This species is common in Palaeocene sediments of the Murray and Great Artesian Basins, and those of the Brisbane area. It differs from Teneridinium magnoides Krutzsch, 1962 in being much smaller and with less prominent apical and antapical horns. S. hansonianum has a more pronounced girdle and antapical horns. Saeptodinium tasmaniensis sp. no v. (Plate 1, Fig. 12; Plate 2, Figs 1-6) Diagnosis: Cavate cysts of ovoidal to slightly peridinioid form. Endocorpus ovoidal enclosed by the pericorpus which has a weakly formed apical horn and two very small antapical bulges. Periphragm scabrate. Endophragm uniformly granulate. Tabulation not marked by ornament. Girdle indistinct or not marked. Archeopyle not observed. Apex provided with a thickened plug. Holotype: Plate 2, Figure 1. Mt Bischoff, Tasmania, Late Oligocene—Early Miocene. Description: Epitract rounded triangular in outline, and hypotract rounded or with two rather sharp antapical bulges separated by a wide indentation 7-10/* deep. Folding often obscures the antapical bulges giving either the impression of a rounded hypotract or one with a single antapical prominence. Pericoel variable, 2-5/* wide and widest in the antapical and apical regions. Mostly the periphragm lies close to the endophragm. Periphragm and endophragm of approximately equal thickness, 0.5-1/*. Archeopyle formed possibly by splitting between hypo- and epitracts. Dimensions: (10 specimens) Overall length 45-62/*. Overall breadth 42-58/*. Remarks: This cyst type resembles S. gravattensis but is distinct in its ornament and shape of the hypotract. Both have a similar apical plug. Known only from the sediments at Mt Bischoff, Tasmania. Genus Morkallacysta nov. Diagnosis: Non-cavate dinoflagellate cysts of strongly peridinioid form with prominent

163 apical and antapical horns. Helicoid girdle strongly marked. Archeopyle triangular with attached operculum. Cyst lacking tabulation and ornament. Type species: Morkallacysta pyramidalis sp. nov. Palaeocene, Victoria. Remarks: The genus superficially resembles Deflandrea but differs in being non-cavate and having an attached operculum.

Morkallacysta pyramidalis sp. nov. (Plate 1, Figs 5-11) Diagnosis: A Morkallacysta of strongly peridinioid form. The epitract is produced into a rounded horn and the antapex into two subequal horns. Cyst smooth to scabrate. Strongly marked and helicoid girdle. Archeopyle triangular, with operculum. Description: Epitract 0.5/* thick, strongly triangular in outline and larger than the hypotract. Antapical bulges variable in degree but usually formed by a deep depression in the mid-line. Girdle about 8/* wide and often deeply depressed, 2-4/* deep, strongly helicoid and laevorotary. Vertical area depressed in the hypotract. Archeopyle large, approximately 30/* wide and 10-12/* high; probably formed by loss of a precingular plate. Operculum attached near margin of the cingulum. Holotype: Plate 1, Figure 5. Renmark Beds, Palaeocene, Victoria. Dimensions: (20 specimens) Length 55-68/* breadth 48-55/*. Remarks: This species is similar to that described from the Princetown Member as Deflandrea obliquipes Deflandre & Cookson, 1955, but differs from that species in lacking an endocorpus. Resemblance to certain cysts of marine Peridinium spp. (Wall & Dale, 1967) is striking. This cyst type is widely distributed in the Murnpeowie Formation of the Great Artesian Basin and the lower Renmark Beds of the Murray Basin. Genus Cubiculosphaera nov. Diagnosis: Cavate dinoflagellate cysts of ovoid form. Two antapical horns weakly or scarcely formed. Pericorpus and endocorpus closely adpressed. Pericorpus similar in shape to endocorpus. Both walls smooth to scabrate lacking tabulation and without trace of cingulum. Archeopyle formation not known. Type species: Cubiculosphaera maslinensis sp. nov. Middle Eocene, South Australia.

Spec.Publs geol.Soc.Aust., 4: pp. 159-166, Pis 1-2, 1973.

WAYNE

164

K.

Remarks: Geiselodinium Krutzsch, 1962 has a better developed apical horn and the type species, G. geiseltalense Krutzsch, 1962 has an archeopyle. Saeptodinium gen. nov. is more strongly peridinioid and the shape of the two walls is dissimilar. Cubiculosphaera maslinensis sp. nov. (Plate 2, Fig. 10) Diagnosis: Endocorpus and pericorpus ovoidal, breadth greater than length. Pericorpus and endocorpus have two slight antapical bulges. Endophragm slightly granulate and periphragm smooth to scabrate. Pericoel very narrow, 1-2/x wide, and endophragm often closely adpressed to periphragm. No girdle or archeopyle observed. Holotype: Plate 2, Figure 10. North Maslin Sands, Middle Eocene, South Australia. Description: This type is distinctive in its shape and is often much folded. Both periphragm and endophragm are of equal thickness, 0.5 -l/x. The fine ornament which is more prominent on the endophragm does not reflect any tabulation. Dimensions'. (10 specimens) Overall length 45-60/*. Overall breadth 53-65/x. Remarks: This species resembles closely in form the extant genus Glenodinium Ehrenberg, 1837. Characteristically the cyst type described here offers little in the way of morphological criteria such as tabulation or archeopyle formation. Known only from the North Maslin Sands clay lens at Maslin Bay, South Australia.

HARRIS

Cleistosphaeridium tenuum sp. nov. (Plate 2, Figs 7-9) Diagnosis: Proximo-chorate cysts of circular or ovoidal shape. Cyst wall two-layered, very thin, the outer layer producing a dense network of short appendages, hollow and distal end anchor-shaped. Archeopyle apical. Girdle not observed. Holotype: Plate 2, Figures 8 & 9. Darra Formation, Palaeocene, Queensland. Description: Cyst more or less circular except for archeopyle region and shape modified by the dense array of appendages. These do not reflect any tabulation and are 4-5/* long, 1.5-2.5/* wide, hollow and branched at their extremities, closed distally and not in communication with endocoel. Branches 2 or 3 in number and 2-3//, long giving an 'anchor' shape to the appendage. Archeopyle is presumably formed by the loss of apical plates and there is some suggestion of a sulcal notch in some specimens. Wall layers less than \jx thick. Dimensions: (10 specimens) Length (without appendages) 22-35/x. Breadth (without appendages) 18-25/*. Remarks: The species is only known from the Brisbane area and reaches a frequency of four per cent of the total sporomorph-microplankton assemblage. The very thin walls and very dense array of short processes distinguish this from other described species.

ACKNOWLEDGMENTS Genus Cleistosphaeridium Davey, Downie, Sarjeant & Williams, 1966 Type species: Cleistosphaeridium diversispinosum Davey, Downie, Sarjeant & Williams, 1966.

My special thanks go to Dr A. R. Loeblich Jr for readily supplying me with a photo-copy of Krutzsch's otherwise unavailable paper. This paper is published with the permission of the Director of Mines, South Australia.

REFERENCES CHURCHILL, D . M . , & SARJEANT, W . A . S.,

1962:

Fossil dinoflagellates and hystrichospheres in Australian freshwater deposits. Nature, Lond., 194, p. 1 0 9 4 . , , 1963: Freshwater microplankton from Flandrian (Holocene) peats of south western Australia. Grana palynol., 3, pp. 29-53.

DAVEY, R . J., DOWNIE, C . , SARJEANT, W . A .

S.,

& WILLIAMS, G. L., 1966: Studies on Mesozoic and Cainozoic dinoflagellate cysts. British Mus. (Nat. Hist.), Bull. Geol., Supplement 3, p. 248. DOWNIE, C . , & SARJEANT, W . A . S.,

1966:

The

morphology, terminology and classification of fossil dinoflagellate cysts. British Mus. (Nat. Hist.), Bull. Geol. Supplement 3, pp. 10-17.

TERTIARY NON-MARINE DINOFLAGELLATE CYST ASSEMBLAGES EVITT, W. R., 1970: Dinoflagellates—a selective review. Geoscience and Man, 1, pp. 29-45. , & DAVIDSON, S. E., 1964: Dinoflagellate

studies, 1. Dinoflagellate cysts and thecae. Stanford Univ. Pubis, Geol. Sci., 10 (1), pp. 1-12. , & WALL, D., 1968: Dinoflagellate studies IV. Theca and cysts of Recent freshwater Peridinium limbatum (Stokes) Lemmermann. Stanford Univ. Pubis, Geol. Sci., 12 (2), pp. 1-15. GOCHT,

H.,

1969:

Formengemeinschaften

66,

pp.

221-227.

HUBER, G . , & NIPKOW, F . ,

1922:

MCGOWRAN, B., HARRIS, W . K . , & LINDSAY, J . M . ,

1970: The Maslin Bay flora, South Australia. 1. Evidence for early Middle Eocene age. Neues Jb. Geol. Paldont. Mh., 1970, pp. 481485. NORRIS, G . , & MCANDREWS, J . H . , 1 9 7 0 : Dinoflagellate cysts from post-glacial lake muds, Minnesota ( U . S . A . ) . Rev. Palaeobot. Palyn., 10, p p . 1 3 1 - 1 5 6 .

Alt-

tertiaren Mikroplanktons aus Bohrproben des Erdolfeldes Meckelfeld bei Hamburg. Palae ontographic a, 126 B, pp. 1-100. HARLAND, R . , 1 9 7 1 : Fossil dinoflagellate cysts from Lake Gnotuk, Victoria, Australia. Proc. R. Soc. Vict., 84, pp. 245-254. , & SARJEANT, W. A. S., 1970: Fossil freshwater microplankton (dinoflagellates and acritarchs) from Flandrian (Holocene) sediments of Victoria and Western Australia. Proc. R. Soc. Vict., 83, pp. 211-234. HARRIS, W. K., 1965: Tertiary microfloras from Brisbane, Queensland. Rep. geol. Surv. Qd, 10. HOUSTON, B . R . , 1 9 6 5 : New and re-defined names in Queensland stratigraphy. Qd Govt Min. J., Experimented

Untersuchungen iiber die Entwicklung von Ceraiium hirundinella O.F.M. Z. Bot., 14, pp. 337-371.

, , 1923: Experimentelle Untersuchungen liber die Entwicklung von Ceratium hirundinella O. Fr. Mull. Flora, n.s., 16, pp. 114-215. KRUTZSCH, W . , 1 9 6 2 : Die Mikroflora der Geiseltalbraunkohle. Teil III, Susswasserdinoflagellaten aus subaquatisch gebildeten Blatterkohlenlagen des mittleren Geiseltales. Hallesches Jb. Mitteldt. Erdgesch, 4, pp. 4045. LANGE, R. T., 1970: The Maslin Bay flora, South Australia. 2. The assemblage of fossils. Neues Jb. Geol. Paldont. Mh., 1970, pp. 486-490.

165

Cainozoic foraminifera and stratigraphy of the Adelaide Plains Sub-basin, South Australia. Bull. geol. Surv. S. Aust., 42.

LINDSAY, J . M . , 1 9 6 9 :

SARJEANT, W . A . S . & STRACHAN, I . , 1 9 6 8 :

Fresh-

water acritarchs in Pleistocene peats from Staffordshire, England. Grana palynol., 8, pp. 204-209. SMITH, G. M., 1950: The Freshwater Algae of the United States (2nd Ed.). McGraw-Hill, New York. STAPLIN, F. L., 1961: Reef-controlled distribution of Devonian microplankton in Alberta. Palaeontology, 4, pp. 393-424. TAYLOR, D . J . , 1 9 6 4 : Foraminifera and the stratigraphy of the western Victorian Cretaceous sediments. Proc. R. Soc. Vict., 77, pp. 535602. TRAVERSE, A., 1955: Pollen analysis of the Brandon lignite of Vermont. Rep. Invest. U.S. Bur. Mines, 5151. WALL, D., 1965: Microplankton and spores from the Lower Jurassic of Britain. Micropaleontology, 11, pp. 151-190. , & DALE, B., 1967: The resting cysts of modern marine dinoflagellates and their palaeontological significance. Rev. Palaeobot. Palyn., 2, pp. 349-354. , & , 1968: Modern dinoflagellate cysts and evolution of the Peridiniales. Micropaleontology, 14, pp. 265-304. , & , 1970: Living hystrichosphaerid dinoflagellate spores from Bermuda and Puerto Rico. Micropaleontology, 16, pp. 47-58. WALL,

Wayne K. Harris, Geological Survey of South Australia, Department of Mines, P.O. Box 38, Rundle Street, Adelaide, South Australia 5000.

Spec.Publs geoLSoc.Aust., 4: pp. 1 5 9 - 1 6 6 , Pis 1 - 2 , 1 9 7 3 .

D.,

GUILLARD,

R.

R.

L.,

&

DALE,

B.,

1967: Marine dinoflagellate cultures from resting spores. Phycologia, 6 (2), pp. 83-86.

WAYNE K. HARRIS

166

EXPLANATION OF PLATES PLATE 1

All Figures X 500 and all but Figures 4 & 5 by Nomarski differential interference contrast. Saeptodinium gravattensis gen. et sp. nov. Figures 1-4 Brisbane NS5, core at 270 ft (82.3 m) Slide No. Catalogue No. Coords 1. S103/4 P 720 47.4, 104.7 2. S103/4 P 721 44.0, 102.7 3. S103/3 P 722 48.9, 110.0 4. S103/4 P 723 35.9, 100.8 Morkallacysta pyramidalis gen. et sp. nov. Figures 5-11 A.A.O. Morkalla No. 1 Well. Sidewall core at 1809 ft (551.4 m) 5. P 724 S2142/2 33.9, 103.5 P 725 95.9 S2142/1 27.1, 6. 7. S2142/1 P 726 38.0, 104.5 S2142/1 8. P 727 45.2, 95.7 S2142/1 9. P 728 31.0, 103.2 S2142/1 P 729 31.8, 107.1 10. S2142/4 P 730 11. 44.5, 107.2 Saeptodinium tasmaniensis gen. et sp. nov. 12. Mt Bischoff, beneath basalt. S1021/4 27.2, 108.7 P 731

Figures 1-8 X 500, 9 & 10 x 1250.

PLATE 2

Figures 1, 4, 5, 7, 8, 10 by Nomarski differential interference contrast. Figure 9 by phase contrast. Saeptodinium tasmaniensis gen. et sp. nov. Figures 1-6 Mt Bischoff, beneath basalt. Coords Slide No. Catalogue No. 43.8, 103.0 S1021/5 P 732 1. 29.8, 99.0 2. S1021/1 P 733 40.2, 95.5 3. S1021/1 P 734 4. P 735 25.9, 106.9 S1021/3 5. 27.1, 102.1 S1021/3 P 736 P 737 6. S1021/1 23.0, 106.2 Cleistosphaeridium tenuum sp. nov. Figures 7-9 Brisbane NS5, core at 270 ft (82.3 m) 7. S103/1 P 738 34.4, 8. S103/4 P 739 46.2, 9. S103/4 P 739 46.2, Figure 10 Maslin Bay clay

105.1 108.5 108.5

Cubiculosphaera maslinensis gen. et sp. nov. S1308/1 P 740 37.1, 105.5

WAYNE K .

HARRIS

Spec.PubIsgeol.Soc.Aust., 4, 1973.

PLATE 1

PLATE

2

WAYNE K .

HARRIS

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA (DINOPHYCEAE) IN VICTORIAN COASTAL AND OFFSHORE BASINS, AUSTRALIA By LEWIS E. STOVER (With 9 Text-Figures and 5 Plates) ABSTRACT

Specimens of the dinophyceate genus Deflandrea occur fairly consistently in Palaeocene and Eocene palynomorph assemblages from the Gippsland, Bass, and Otway Basins, southeastern Australia. From the Latrobe Group and its chronostratigraphic equivalents, twenty species of Deflandrea are identified of which D. conorata, D. druggii, D. extensa, D. flounderensis, D. medcalfii, and D. truncata are proposed as new taxa, and D. granulosa Cookson & Eisenack is regarded as a junior synonym of D. phosphoritica Eisenack. Apart from the descriptions of the new forms, additional morphological and stratigraphic information are given for each of the previously described species. The variability and relationships of various features found among species of Deflandrea from southeastern Australia are discussed; these include the overall shape of the cyst, apical and antapical horn development, amount and clarity of tabulation, archeopyle style, sculpturing, and the position and size of the endoblast relative to the periblast. Consideration of these features individually and in different combinations indicates those characters useful in determining the relative ages of Deflandrea-beaxmg assemblages, and in suggesting trends among Palaeocene and Eocene species. Based on species from the Victorian coastal and offshore basins, some general tendencies among forms with free opercula are: 1, the length to width ratio of the archeopyle decreases with decreasing geological age; 2, the clarity of the tabulation and the prominence of the cingulum and sulcus decrease with decreasing geological age; 3, apical and antapical horn development become subdued with decreasing geological age, although exceptions do occur and concomitantly the outline tends to become more rotund and the endoblast occupies a greater proportion of the pericoel; and 4, coarse sculpturing is confined to species from the younger part of the section.

INTRODUCTION The fossil dinophyceate cyst genus Deflandrea occurs fairly consistently in nearshore Palaeocene and Eocene palynomorph assemblages from sedimentary sections in the coastal and offshore basins of Victoria. Since the genus was first described from the Oligocene of Germany (Eisenack, 1938) numerous Late Cretaceous and Tertiary species have been reported from every continent; thus, Deflandrea has a wide geographic distribution. Deflandre & Cookson (1955) reported the first occurrence of the genus in Australia and, in the same publication, they erected Deflandrea bakerii, D. bakerii f . pellucida, D. heterophlycta, D. obliquipes, D. pachyceros and D. robusta. Ten years later Cookson & Eisenack (1965a, b, c) described D. delineata, D. dilwynensis, and D. pentaradiata from Palaeocene rocks, and D. dartmooria, D. granulosa and D. leptodermata from Eocene strata.

Cookson & Eisenack also identified D. phosphoritica in Victoria. This species was reported earlier from Western Australia (Cookson & Eisenack, 1961). Purposes of this report are to provide additional information on previously described species, to introduce six new species, to discuss the morphological variability and trends among Palaeocene and Eocene species of Deflandrea, and lastly to comment on the stratigraphic utility of the genus in southeastern Australia. Material. More than 100 samples, all containing specimens of Deflandrea, were processed and examined. Of these, about 30 were collected from the Palaeocene and Eocene coastal sections between Princetown and Anglesea, Victoria, and include topotypic material from the Pebble Point Formation, the Dilwyn Formation, the Johanna River Sands and the Browns Creek Clays. Approximately

Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

LEWIS E. STOVER

168

80 conventional and sidewall core samples from offshore wells in the Bass and Gippsland Basins were used in this study. Through the courtesy of the National Museum of Victoria, I examined all of the type specimens of Deflandrea in their collection and recorded the associated microplankton and spore-pollen assemblages in the strew preparations. Several localities from which Deflandre & Cookson (1955) and Cookson & Eisenack (1965 a, b, c) obtained Palaeocene and Eocene samples are along the southwestern Victorian coast from Castle Cove to Princetown. The stratigraphy of this area was presented succinctly by O. P. Singleton (1967) and the Palaeocene and Early Eocene spore-pollen assemblages from sections between Dilwyn Cove and Princetown were described and illustrated by W. K. Harris (1965). Earlier this year (1971) D. J. Taylor and I collected samples from most of those localities from which Dr I. C. Cookson and her co-workers designated type specimens. These localities and the geographic coordinates of Bass and Gippsland Basin wells from which type specimens were chosen for the taxa proposed in this paper are listed below. Otway Basin: Locality 1. Section near the mouth of the Gellibrand River about 1.2 km southeast of Point Ronald; Victoria Princetown-Moonlight head district; locality (c) of Deflandre & Cookson (1955, p. 246). Locality 2. Section between Point Ronald and Point Margaret about 2.1 km southeast of the mouth of the Gellibrand River; Rivernook locality in Cookson & Eisenack (1967, p. 247). Locality 3. Section between Point Ronald and Point Margaret at about 2.4 km southeast of the mouth of the Gellibrand River, Dilwyn Clay 54 m above the base of the Pebble Point Formation. Locality 4. Section at Point Margaret, about 2.7 km southeast of the mouth of the Gellibrand River, Pebble Point Formation at 12.3 to 14.3 m above the base of the formation. Locality 5. Northwest side of the Dilwyn Cove about 4.2 km southeast of the mouth of the Gellibrand River; Pebble Point sample approximately 9.1 m above the base of the formation, Cookson & Eisenack (1965c, p. 139). Locality 6. Southeast side of Dilwyn Cove about 4.8 km southeast of the mouth of the Gellibrand River; Pebble Point samples between 1.3 and 2.0 m above the base of the formation, Cookson & Eisenack (1965c, p. 139). Locality 7. Outwash gully section about 0.5 km southeast of the mouth of the Johanna River; Browns Creek Clays locality in Cookson & Eisenack (1965a, p. 120). Locality 8. Coastal section at Castle Cove about 3.3 km east southeast of the mouth of the Johanna River; Castle Cove, Aire coast locality in Deflandre & Cookson (1955, p. 247). Bass Basin: Locality 9. Bass-1, Locality 10. Bass-2, Gippsland Basin: Locality 11. Albacore-1, Locality 12. Barracouta-3, Locality 13. Dolphin-1, Locality 14. Flounder-1, Locality 15. Flounder-2, Locality 16. Flounder-3, Locality 17. Groper-1, Locality 18. Marlin-1, Locality 19. Perch-1, Locality 20. Tuna-3, Locality 21. Turrum-1, Locality 22. Glencoe-4,

Latitude (S)

Longitude (E)

39° 46' 18" 39° 53' 09"

145° 44' 03" 146° 18' 18"

38° 34' 00" 38° 19' 19" 38° 29' 32" 38° 18' 52" 38° 19' 18" 38° 18' 58" 38° 56' 20" 38° 14' 03" 38° 14' 37" 38° 10' 10" 38° 12' 10" 38° 11' 00"

148° 19' 54" 147° 37' 05" 147° 22' 43" 148° 25' 29" 148° 26' 53" 148° 28' 23" 147° 24' 56" 148° 15' 33" 147° 19' 24" 148° 10' 10" 148° 14' 41" 147° 05' 00"

DEFLANDREA MORPHOLOGY Most of the morphologic features on fossil cysts of Deflandrea are depicted in the generalized drawing shown in Figure 1. The cyst or tract is composed of an inner body, endoblast, made up of the endophragm or wall and the enclosed space, the endocoel. Surrounding the endoblast is the periblast, with its wall or periphragm. The space between the periphragm and the endophragm is termed the pericoel. Commonly, the anterior and posterior parts of the periphragm are attenuated to form the apical and antapical horns, respectively. A transverse depression, the cingulum, divides the cyst into an anterior part, the epitract, and a posterior part, the hypotract. On the ventral surface a broad, shallow, longitudinal depression, the sulcus, extends posteriorly from near the separated ends of the cingulum. The dorsal and ventral surfaces of the periphragm may be divided into polygonal fields or plates, whose number and arrangement define the tabulation. The plates are organized in rows with the plates in a given row constituting a series. The series, because of their position on the periphragm, are named beginning at the apical end, the apical, precingular, cingular, postcingular and antapical series. An intermediate series, the anterior intercalary, lies between the apical and precingular series on the dorsal surface. The individual plates may be designated by a number, indicating its sequential number in a row, and an exponential symbol, indicating to which series it belongs. Thus the first plate of the apical series may be signified by 1', the third by 3' and the fourth plate of the postcingular series by 4"\ Other designates are given in Figure 1 which also indicates the basic tabulation for Deflandrea as 4', 3a 7" 5c 5"', 2"". On a few species, irregularly shaped, small sulcal plates are identifiable in the sulcus and on a majority of species the cingulum is undivided. An opening on the anterior dorsal surface of the periphragm, the intercalary archeopyle, occurs on all species of Deflandrea, and its cover or operculum may not be in place. A corresponding opening usually occurs in the endophragm. DESCRIPTIVE PALAEONTOLOGY Species of Deflandrea are treated in the following order, which, in general, corresponds also to their arrangement on the Plates. D. bakerii D. flounderensis D. pellucida D. medcalfii

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA 169 Material: Morphological features on holoD. dartmooria type are obscure owing to desiccation of the D. truncata preparation. Because of the poor condition of D. leptodermata the holotype, Dr Cookson provided 2 slides D. phosphoritica (P23080 and P23081) which contain 4 speciD. granulosa mens of Defiandrea bakerii. Another specimen, D. heterophlycta in the preparation with the holotype, is slightly D. extensa D. sp. cf. D. diebelii smaller but otherwise identical with the holotype insofar as could be ascertained. Three more specimens were identified in the slide SYSTEMATIC DESCRIPTIONS containing the holotype of Eisenackia crassiClass DINOPHYCEAE Pascher, 1914 tabulata Deflandre & Cookson, 1955; and a Order PERIDINIALES Haeckel, 1894 sample from the Pebble Point Formation colFamily DEFLANDREACEAE Eisenack, 1938 lected 1.3-1.5 m above the base of the formaGenus Defiandrea Eisenack, 1938 tion at Point Margaret, Victoria, provided a few well-preserved specimens. The revised Type species: Defiandrea phosphoritica description is based on the dozen or so comEisenack, 1938, p. 187, Text-Figure 6. Defiandrea bakerii Deflandre & Cookson, 1955 plete specimens mentioned above. Revised description: In dorso-ventral view, (Plate 1, Figs 1, 2) lateral margins of the periphragm are 1955 Defiandrea bakerii Deflandre & Cookson, the more or less evenly and gently convex along p. 251, Plate 4, Figures 1, 2, 4. most their length with no apparent demar1960 Defiandrea bakerii Deflandre & Cookson; cationofbetween the epitract and hypotract. A Douglas, p. 19, Plate 3, Figures 18a, slight inflection occurs along the lateral 18b. margins anterior to the archeopyle in 1970 Defiandrea bakerii Deflandre & Cookson; some specimens. Above the inflection, the Deflandre et al., Plate 4, Figures 1, 2, 4, margins of the periphragm converge to form and supplementary Plate 1, Figures 5, 6. a short, usually blunt apical horn. Antapical D. druggii D. conorata D. dilwynensis D. pentaradiata D. robusta D. pachyceros D. obliquipes D. delineata

OPERCULUM PERIPHRAGM / PERICOEL ARCHEOPYLE

APICAL HORN 2'

PERIBLAST (WALL AND CAVITY)

PLATES

ENDOBLAST (WALL AND CAVITY) CINGULUM SULCUS

END0C0EL ENDOPHRAGM

DORSAL SURFACE

VENTRAL SURFACE

Fig. 1. Morphological features of Defiandrea. Spec.Publs geol.Soc.Aust, 4: pp. 167-188, Pis 1-5, 1973.

ANTAPICAL HORNS

LEWIS E. STOVER

170

margin is straight or nearly so on most specimens and truncates the lateral margins rather abruptly. On one specimen (Plate 1, Fig. 1), the antapical margin has a medial peak between shallow concavities. A small, hollow projection about 2^ long may occur at one antapical corner, or a broadly rounded right antapical horn may be developed. Endophragm is circular to subcircular in dorso-ventral view, folded in nearly all specimens, and lies approximately midway between the apical and antapical ends of the periblast. Lateral margins of the endophragm are close to or in contact with the inner surface of the periphragm. Endophragm is smooth, about 1/x thick; periphragm varies in thickness from slightly less than 1 to nearly 2/x. Periphragm surface is weakly verrucate (Plate 1, Fig. 2); verrucae low, irregularly polygonal and separated by narrow grooves. Specimens show no definitive indications of cingulum or sulci. Archeopyle is intercalary, hexagonal, greater in height than in width and the operculum is attached, hinged along its antapical margin. Cracks and tears in the endophragm suggest an opening most likely occurs in this body, at least on some specimens. Measurements for the holotype are: periblast, 112 x 78/x; endoblast, 70 x 70/a. The length of the periblast on other specimens is 93 to 130^,, and the width is 63 to 80p. Types'. Holotype, specimen ilustrated by Deflandre & Cookson (1955, Plate 4, Fig. 1). Paratypes, in the same publication (Plate 4, Figs 2, 4 ) . Type locality: Locality 5. Type strata: Pebble Point Formation, Lygistepollenites balmei spore-pollen Zone, Middle Palaeocene. (The spore-pollen zone names used in the systematic description are those proposed by Stover & Evans elsewhere in this volume). Exact position within the formation of the sample with the holotype is unknown. Additional occurrences'. Specimens of Deflandrea bakerii have been recovered from samples of basal Pebble Point Formation at Point Margaret, Victoria; Douglas (1960) recorded D. bakerii from the Haywood-10 bore in southwestern Victoria and Wilson (1967) reported the species from the Teurian (Palaeocene) Garden Cove Formation, Campbell Island.

Deflandrea pellucida Cookson & Eisenack, 1958 (Plate 1, Figs 7a, 7b) 1955 Deflandrea pellucida forma pellucida Deflandre & Cookson, p. 251, Plate 4, Figure 3. 1958 Deflandrea pellucida Cookson & Eisenack, p. 27, Plate 4, Figure 9. 1960 Deflandrea pellucida Cookson & Eisenack; Douglas, p. 19, Plate 3, Figures 15a, 15b, 17a, 17b. Material: The holotype and 5 unillustrated specimens in the preparation containing the holotype, and illustrations in Douglas (1960). Revised description: In dorso-ventral view the circular to subcircular endoblast is centred or nearly centred along the longitudinal and transverse axes of the specimens. Epitractal part of periphragm is convexly triangular and has a short, narrowly to moderately rounded apical horn; the hypotractal part has convexly arcuate lateral margins, as on the holotype, or the right lateral margin is less convex than the left lateral margin. On some specimens a small, nipple-like projection occurs at the tip of the right antapical horn. Endoblast is large relative to the size of the periphragm and for approximately onethird the length of the specimen, the periphragm and endophragm are in contact laterally; thus the apical and antapical parts of the pericoel are clearly separated. Periphragm and endophragm are each ca 1/x thick; the latter is smooth, or a roughened or finely granular sculpture is discernible on some specimens. The periphragm has low, small grana which are generally confined to the areas beyond the margins of the endoblast. On some specimens there are areas on the periphragm on which the grana are aligned in short rows that might be traces of tabulation. Most specimens show no indication of a cingulum and the position of the longitudinal sulcus is at best poorly defined by subparallel folds on the ventral hypotractal surface. The intercalary archeopyle is hexagonal, slightly wider than long, and the operculum may be free or hinged along its anterior margin. No definite opening was observed in the endophragm. Aside from minor variations in the outline of the periphragm, little variability was seen among the few specimens of Deflandrea pellucida recorded thus far. Specimens are 115-130^ long and 82-90/x wide; endoblast is 70-75/x long.

PALAEOCENE AND EOCENE SPECIES OF Comparison: Deflandrea pellucida differs from D. bakerii by having a less elongate outline and, in general, possessing more pronounced apical and antapical horns, although the horns are not well developed in either species. The archeopyle on D. pellucida is less elongate apically and proportionately wider than on D. bakerii and the sculpturing on the periphragm is coarser in D. bakerii. Types: Holotype, specimen illustrated by Deflandre & Cookson (1955, Plate 4, Fig. 3); no paratype. Type Locality: Nelson bore (^Glenelg No. 1 in Douglas, 1960) in extreme southwestern Victoria. Type stratum: Early Tertiary rocks in the Nelson bore at 1181.5 m, probably Middle Palaeocene. Associated spore-pollen species indicate the holotype is from the lower part of the Lygistepollenites balmei Zone. Additional occurrences'. Deflandre & Cookson (1955) reported the species from the Pebble Point Formation in Victoria. Deflandrea druggii sp. no v. (Plate 1, Figs 3a, 3b, 4) Material: Eighteen specimens from conventional core samples from Flounder-2 and Flounder-3 wells. Description: Outline of periphragm in dorso-ventral view is broadly elliptical, rarely elongate and with a shallow antapical concavity which separates poorly developed antapical horns. Apical margin may be evenly rounded or a short, broadly-based, blunt apical horn may be present. Periphragm is about thick, and is smooth, scabrate or irregularly granulate with the sculptural features varying in diameter from ca 0.5/x to 2.0^. Usually the coarser sculpturing is concentrated on the areas beyond the limits of the endoblast. Surface of periphragm lacks traces of tabulation except for small intercalary archeopyle. Endoblast outline circular or nearly so in dorso-ventral view, commonly modified by folding; endoblast occupies a major part of the pericoel, and is not in contact laterally with the inner surface of the periphragm. Endophragm is ca thick, smooth or faintly scabrate. No opening observed in the endophragm in a position corresponding to the archeopyle in the periphragm. Operculum is relatively small, usually with a narrow apical margin, oblique and slightly convex sides,

DEFLANDREA

171

rounded antapical corners and is hinged antapically. Specimens vary in width (92-103/*) and length (104-122fx); length:width ratio lies between 1:0.83 and 1:0.93. In nearly all specimens in which folding of the endoblast is not severe, its width exceeds its height. Comparison: Specimens of Deflandrea druggii resemble those of D. cretacea Cookson, 1956 in outline, by having the endoblast filling a comparatively large part of the pericoel and by lacking prominent apical and antapical horns. The new species differs by being about twice the size of D. cretacea and by having the endoblast consistently separated laterally, as well as apically and antapically, from the periphragm. Of the specimens illustrated and assigned to D. cretacea from California, U.S.A. (Drugg, 1967) the example shown on his Plate 2, Figure 17 is probably conspecific with D. druggii. Specimens of D. cretacea from the Nelson bore in southwestern Victoria (Cookson 1956, Plate 1, Figs 1-5; not Figs 6 and 7) have the endoblast touching the periphragm laterally, except for her Figure 5 in which the two bodies of the cyst are clearly separated laterally. In this respect and also by having the right antapical horn somewhat better developed, the specimen (Cookson's Fig. 5) resembles more closely those of D. druggii than do her other specimens of D. cretacea. Types: Holotype (Plate 1, Fig. 3) and paratype (Plate 1, Fig. 4 ) . Type locality: Locality 15. Type strata: Latrobe Group, 2,530.32,531.2 m, Early to Middle Palaeocene, Tricolpites longus Zone. Additional occurrence: Flounder-3 (Loc. 16) 2,548.0-2,549.0 m. Deflandrea conorata sp. nov. (Plate 1, Figs 8a, 8b ) Material: Thirty-six specimens. Description: Outline in dorso-ventral view is somewhat lozenge-shaped with slightly convex and rounded lateral margins. Apical horn is short, usually blunt, weakly to moderately well developed, not prominent; antapical horns are rounded, fairly well differentiated, and unequal. On some specimens the larger antapical horn is more pointed than the smaller horn. Periphragm is ca 1/x thick, faintly and uniformly scabrate to finely granulate. Traces of tabulation on the periphragm are lacking

Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

172

LEWIS E. STOVER

except for the intercalary archeopyle and a poorly defined cingulum indicated by faint, parallel, discontinuous transverse linear markings, ca 5^ apart, and located at the widest part of the cysts. Endoblast is circular to elliptical in dorsoventral view, smooth to faintly scabrate, from less than 1 to thick, and commonly folded. The endoblast occupies a major part of the pericoel and the margins of the endophragm do not touch the inner surface of the periphragm laterally. Comparatively small rectangular trapezoidal opercula, with generally rounded corners and apparently hinged antapically, occur on all specimens. Only a few specimens indicate an opening in the endophragm. Specimens of Deflandrea conorata vary in length (140-162/1) and width (98-124/*). Length : width ratio lies between 1:0.8 and 1:0.7. Endoblast is 92-112/* long and 85-114/x wide. The length is greater than the width on some specimens whereas on others the reverse is true; but reliable dimensions of the endoblast are difficult to obtain because of folding. Measurements are of 10 similarly oriented specimens. Comparison: Specimens of Deflandrea conorata are similar to those of D. pellucida and differ from the latter by having a larger endoblast, relative to the periblast, and a rectangular to trapezoidal archeopyle whose height is at least equal to its width and generally greater. The sculpturing on the periphragm is also more evenly distributed on D. conorata in contrast to its interrupted distribution on D. pellucida. Types: Holotype (Plate 1, Fig. 8); an additional 9 specimens are in the preparation with the holotype. Type locality: Locality 16. Type stratum: Latrobe Group, 2,552.52,553.5 m, Early to Middle Palaeocene, Tricolpites longus Zone. Additional occurrences: Flounder-2 (Loc. 15), 2,530.8—2,531.2 m. Deflandrea dilwynensis Cookson & Eisenack, 1965 (Plate 1, Figs 5, 6a, 6b) 1965 Deflandrea dilwynensis Cookson & Eisenack, p. 141, Plate 18, Figures 6-9 [1965c]. Material: Approximately 40 specimens including the holotype and paratypes.

Comments: The general characteristics and diagnostic features for Deflandrea dilwynensis are shown quite well in the photomicrographs of the holotype and paratypes (Cookson & Eisenack, 1965c). Study of additional specimens and the type material indicates that the endophragm is smooth, ca 1/x thick. The intercalary archeopyle is large relative to the size of the epitract and the operculum remains attached. Not all specimens show the archeopyle and on those that do the apical and lateral margins are sharply defined whereas its antapical margin is undefined or marked by only a faint line on the periphragm. Splits leading from the basal corners of the archeopyle to the cingulum tend to develop. As a result the archeopyle may be misinterpreted as being precingular rather than intercalary. An opening in the endophragm appears to approximate the archeopyle in size and shape: however, an endophragmal opening was observed on only a few specimens. On these, the inner opercula are apparently attached antapically. Types: Holotype, specimen illustrated by Cookson & Eisenack (1965c, Plate 18, Figs 7, 8); paratypes, specimens illustrated in the same publication (Plate 18, Figs 6, 9). Type locality: Locality 5. Type stratum: Pebble Point Formation, about 9.1 m above the base of the formation, Middle Palaeocene. Lygistepollenites balmei Zone. Deflandrea pentaradiata Cookson & Eisenack, 1965 (Plate 2, Fig. 6) 1965 Deflandrea pentaradiata Cookson & Eisenack, p. 139-140, Plate 18, Figures 1, 2[1965c]. Material: Holotype, paratype, and seven additional specimens from the type locality. Comments: The cingular projections and sharply pointed, clearly separated antapical horns are characteristic of the species. Periphragm and endophragm are each ca 1.5/x thick; however, the measurements are somewhat misleading because of surface irregularities. These irregularities impart a finely striate pattern that is just discernible over most of the periphragm and is shown best adjacent to the cingulum. Surface of the endophragm is smooth. The archeopyle is trapezoidal with a broad base and narrow apical end and is longer than wide; its operculum consists of

PALAEOCENE AND EOCENE SPECIES OF two closely appressed pieces with the piece from the endophragm being shorter than that from the periphragm. Types'. Holotype, specimen illustrated by Cookson & Eisenack (1965, Plate 18, Fig. 1). Paratype illustrated in the same publication (Plate 18, Fig. 2 ) . Type locality: Locality 6. Type stratum: Pebble Point Formation, 1.3 m above the base of the formation, Middle Palaeocene Lygistepollenites balmei Zone. Deflandrea

robusta Deflandre & Cookson, 1955 (Plate 2, Fig. 5) 1955 Deflandrea robusta Deflandre & Cookson, p. 250, Plate 4, Figure 9. Material: Species is known f r o m a single specimen only. Comments on the holotype: The periphragm is transparent, 1 to 1.5^ thick and has a short apical horn, ca 15/x in height, with a broad base and a slightly thickened tip. Antapical horns are subequal with narrowly rounded tips. Along the lateral margins the position of the cingulum is marked by minute projections on one side and a gentle thickening of the periphragm on the other. Trace of the cingulum is faint on the dorsal and ventral surfaces. Small widely spaced grana occur on the epitract and there is a slight concentration of grana on the apical horn. Endophragm is about 1/x thick and faintly scabrate. The intercalary archeopyle is hexagonal, relatively large (45/z wide, 23^ long), and the operculum is lacking. There is some indication that an opening occurs also in the endophragm; however, its shape could not be determined because of folding. Cyst is 133/x, long and 84ja wide at the cingulum; endophragm is 77^ wide. Type: Holotype, specimen illustrated by Deflandre & Cookson (1955, Plate 4, Fig. 9 ) . No paratypes. Type locality: Locality 1. Type strata: About 2-2.5 m of dark grey to black carbonaceous siltstone of the upper part of the Dilwyn Formation. Exact position of sample from which the holotype was obtained is unknown. Associated abundant, diverse, and well preserved spores and pollen indicative of the Early Eocene Malvacipollis diversus Zone occur in the preparation with the holotype.

Deflandrea

DEFLANDREA

173

pachyceros Deflandre & Cookson, 1955

(Plate 2, Figs la, lb, 2, 3a, 3b) 1955 Deflandrea pachyceros Deflandre & Cookson, p. 252, Plate 4, Figure 7. Material: Holotype, plus 35 additional specimens. Comments: Deflandrea pachyceros is quite similar to D. obliquipes except that in the former the horns are shorter, the sculpturing tends to be coarser and the overall size of the specimens is less (Plate 2, Fig. 3). However, specimens showing characteristics intermediate between the two species have been observed and it may be that D. pachyceros and D. obliquipes represent the end members of a rather variable species. It is also possible that the species are distinct and only certain features on each overlap. In either case, more specimens are needed before a critical analysis of the two species is made. On most specimens of D. pachyceros, longitudinal splits occur on the dorsal surface of the periphragm between the cingulum and the basal corners of the archeopyle. These are interpreted as coinciding with the lateral borders of plate 4"; on some specimens accessory splits, also anterior to the cingulum, occur near the lateral margins of the periphragm; these are interpreted as the boundaries between plates 2" and 3" and between 5" and 6". The opening in the endophragm is seen in relatively few specimens (Plate 2, Fig. 1). The shape and size of this opening, like those in the archeopyle, are difficult to ascertain because of folding and splitting of the walls. Types: Holotype, specimen illustrated by Deflandre & Cookson (1955, Plate 4, Fig. 6). N o paratypes. Type locality: Locality 1. Type strata: Same as for Deflandrea obliquipes, Early Eocene Malvacipollis diversus Zone. Additional occurrences: Specimens have been obtained from the Dilwyn Formation, from approximately 50 m above its base to the top of the formation in the Princetown area, Victoria; from Bass-2 well (Loc. 10) at 1,626.4 m; and f r o m Marlin-1 well (Loc. 18), at 1,545.1-1,556.7 m, Gippsland Basin. Deflandrea

Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

obliquipes Deflandre & Cookson, 1955 (Plate 2, Figs 7, 8a-c)

174

LEWIS E. STOVER

1955 Defiandrea obliquipes Deflandre & Cookson, p. 252, Plate 4, Figure 6. 1967 Defiandrea obliquipes Deflandre & Cookson, Cookson & Eisenack, p. 248, Plate 39, Figures 9, 10. Material: Eighteen specimens, including the holotype. Comments: The asymmetrical endoblast is apparently characteristic of the species and in nearly all of the specimens the apical margin and the antapical corners of the endophragm protrude into the bases of the horns. Sculpturing on the periphragm consists of irregularly shaped grana (Plate 2, Figs 7, 8c) which tend to be densest on the surfaces overlying the endoblast and become less dense and even smaller on the horns. Also, the cingulum is either less sculptured than the adjacent areas or is smooth. Archeopyle margins are poorly defined and commonly the dorsal precingular plates are incompletely split apart so that the shape and dimensions of the opening are uncertain. Types: Holotype, specimen illustrated by Deflandre & Cookson (1955, Plate 4, Fig. 6). No paratypes. Type locality: Locality 1. Type strata: About 2-2.5 m of dark grey carbonaceous siltstone from the upper part of the Dilwyn Formation. Exact position of sample from which the holotype was obtained is unknown. Section contains spores and pollen indicative of the Early Eocene Malvacipollis diversus Zone. Additional occurrences: Defiandrea obliquipes occurs sporadically in the Dilwyn Formation from approximately 50 m above the base to the top of the formation in the Princetown area, Victoria. Although Deflandre & Cookson (1955) reported the species from the Pebble Point Formation, no specimens were illustrated from this formation; numerous preparations from the Pebble Point Formation and its equivalents have failed to reveal specimens of D. obliquipes. Defiandrea delineata Cookson & Eisenack, 1965 (Plate 2, Fig. 4) 1965 Defiandrea delineata Cookson & Eisenack, pp. 140-141, Plate 18, Figures 3-5, Text-Fig. 1 [1965c]. Material: Holotype and paratype. Comments: Indications of tabulation are limited to the areas of the precingular and

postcingular plates and are clearer on the holotype than on the paratype. Sculpturing on the periphragm consists of low, short, discontinuous wavy ridges interspersed with small grana (Plate 2, Fig. 4). The periphragm is ca 1.5^ thick, but appears thicker owing to the compressed condition of the type specimens. Contrary to the statements by Cookson & Eisenack (1965c), the archeopyle is at least as long as it is wide (not wider than long) and the cingulum does not pass over the boundaries of the sulcus, but terminates at its margins. Types: Holotype, specimen illustrated by Cookson & Eisenack (1965c, Plate 18, Figs 4, 5). Paratype, illustrated in the same publication (1965c, Plate 18, Fig. 3). Type locality: Locality 6. Type stratum: Pebble Point Formation, 1.3 m above the base of the formation, Middle Palaeocene Lygistepollenites balmei Zone. Defiandrea flounderensis sp. nov. (Plate 3, Figs la-c, 2) Material: More than 100 specimens. Description: Periphragm outline triangularly elongate in dorso-ventral view. Apical horn usually prominent and well defined; antapical horns less well defined and with the right horn commonly longer than the left, but not greatly so. On many specimens, the dorsal antapical part of the periphragm forms a skirt-like membrane between the horns (Plate 3, Figs 1, 2) with a nearly straight antapical margin; however, in the same area on the ventral surface the margin of the periphragm is concave between the antapical horns. Lateral margins convex to straight and interrupted at about midheight by the ridges of the cingulum. Periphragm is ca 1 /x thick, in part smooth and in part finely granulate or conate with the sculpturing corresponding to plate areas. Tabulation peridinioid (4', 3a?, 7", 5'", 2""). On most specimens the shapes and boundaries of the apical plates, except of 1', and the lateral plates are difficult to determine; whereas those for the medial plates are usually evident. Cingulum is marked by parallel transverse ridges, commonly with beaded crests, offset ventrally, undivided, and the floor of the cingulum is smooth, or rarely with scattered grana. Sulcus generally poorly defined, and a comma-shaped scar is discernible on some specimens in the sulcus near its junction with the cingulum. In dorso-ventral view, the outline of the

PALAEOCENE AND EOCE* ' SPECIES OF DEFLANDREA endoblast is circular or nearly so. Endophragm is smooth, granulate, or coarsely vermiculate and 1-2.5/x thick. Almost invariably the endophragm is thicker than the periphragm. Laterally, the periphragm and endophragm are close together and the distance between the two walls increases apically and antapically. The large intercalary archeopyle is broadly trapezoidal, about as wide as high, and with a relatively narrow apical end. The corresponding opening in the endophragm is as wide as in the periphragm, but the height is only about half as much. Operculum free. Complete specimens are 112-166^ long and 76-98//, wide at the cingulum. Approximately three-quarters of the specimens are less than 135/x long. The length to width ratio is 1:0.551:0.70 (mean 1:0.65). Maximum diameter of the endoblast is 60-88/x and in about threequarters of the specimens, its width is greater than its length. Width of endophragmal opening is 37-46/x and its length is 17-24,* (width consistently about twice the length). Measurements based on 25 specimens. Comparison: Deflandrea flounder ensis differs from D. dartmooria Cookson & Eisenack, 19656 by having a larger length to width ratio, generally more subdued sculpturing on the periphragm, sculpturing on the endophragm (on most specimens), and by lacking linear markings at plate boundaries. Types: Holotype (Plate 3, Fig. 1); paratype (Plate 3, Fig. 2). Type locality: Locality 14. Type stratum: Flounder Formation at 1,966 m, Early Eocene Malvacipollis diversus Zone. Additional occurrences'. Flounder-1 well (Loc. 14) at 1,958.1 m; Flounder-2 well (Loc. 15), at 2,124.0 m-2,381.4 m; Flounder-3 well (Loc. 16) at 2,023.3-2,199.1 m. Deflandrea medcalfii sp. nov. (Plate 3, Figs 3a-c) 1967 Deflandrea dartmooria Cookson & Eisenack p. 248, Plate 39, Figures 7, 8 (not Deflandrea dartmooria Cookson & Eisenack 19656, p. 133, 132, Plate 16, Figs 1, 2, Text-Fig. 1). Material: Approximately 50 specimens. Description: Cysts are tricornate with an elongate, tapered apical horn and prominent antapical horns of about equal size and with narrowly rounded or pointed tips. Antapical horns separated by a broad medial concavity.

175

Lateral margins of periphragm are gently convex anterior to the cingulum, and straight to slightly concave posterior to the cingulum. Periphragm less than thick and clear except for areas with small coni. The species has a typical peridinioid tabulation of 4', 3 a?, 7", 5"', 2"" with the size, shape, and position of the plates represented by the conate areas. Cingulum defined by transverse parallel ridges about high, 5-7/x apart, and fringed with coni, denticles, or spinules. On some specimens the cingulum is subdivided by longitudinal ridges and its floor is smooth or bears scattered coni. Sulcal margins generally indefinite, and on most specimens, a hook-shaped scar occurs in the sulcus near its junction with the cingulum and generally a little right of centre. Endoblast is comparatively large, broadly elliptical to ovoid in dorso-ventral view, generally wider than long although on some specimens the reverse is true, and located about centrally with respect to the periphragm. Endoblast configuration approximates that of the periblast laterally but deviates apically and antapically. Endophragm is smooth and ca 0.5^ thick. Archeopyle intercalary, large, irregularly hexagonal in outline, and with a narrower apical than antapical margin. Opening in endophragm corresponds with that in the periphragm. Operculum free. Length of periphragm is 106-140/x, width of periphragm is 68-75,* and the specimens have a length : width ratio between 1 : 0.53 and 1 : 0.73 (mean 1:0.64). Length of endophragm is 53-70/x, width 58-76^. Apical horn is 22-34,* long; antapical horns 20-36,* long. Measurements based on 12 complete specimens. Comments: Plates 1', 2a, 1", 5" 7", 1"', 3"' and 5"' are usually clearly recognizable. Other plates are displayed with varying degrees of clarity depending upon the density and size of the coni and the orientation of the specimens. Plates on the epitract are nearly always shown better than those of the hypotract. On the apical horn, and to a lesser extent on the antapical horns, the coni tend to be so closely spaced that the horns have a scaly appearance. Comparison: Deflandrea medcalfii differs from D. flounderensis by having a smooth and thinner endophragm, and by having the antapical horns distinctly separated. Generally the apical horn is longer and narrower in D.

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LEWIS E. STOVER

medcalfii and the tabulation is expressed somewhat more clearly. Types: Holotype (Plate 3, Figs 3a-c); paratypes not illustrated. Type locality: Locality 4. Type stratum: Pebble Point Formation, 9.19.5m above the base of the formation, Middle Palaeocene Lygistepollenites balmei Zone. Additional occurrences: In the coastal exposures between Point Margaret and Rivernook, 2.0-2.4 km southeast of the mouth of the Gellibrand River, Deflandrea medcalfii occurs sporadically in the lower part of the Dilwyn Formation up to and including the Rivernook bed. Specimens were also identified from a sidewall core at 2,795.0 m from Albacore-1 well (Loc. 11). Deflandrea

dartmooria Cookson & Eisenack, 1965 (Plate 3, Figs 4a, 4b; Plate 4, Figs la-d, 2) 1965 Deflandrea dartmooria Cookson & Eisenack, pp. 132-133, Plate 16, Figures 1, 2, Text-Figure 1 [19656]. Material: Eight specimens including the holotype; 4 specimens f r o m the Dartmoor Formation; and 3 incomplete specimens from Tuna-3 (Loc. 20). Comments: Outstanding among specimens of Deflandrea dartmooria is the preservation of the complete peridinioid tabulation, even to the extent that several of the minute sulcal plates are recognizable. Individual plates are indicated by concentrations of coni separated

by unsculptured areas. Within the latter are faint linear markings, interpreted as representing plate boundaries. Tabulation is 4', 3a, 7", 5c, 5'", 2"" with probably 7 sulcal plates. Figure 2 depicts the pattern of sulcal plates. Of these, the anterior plates are clearly delimited, but I am uncertain of the plate boundaries in the posterior part of the sulcus; there may be one large posterior plate or perhaps 2 small plates. Some sulcal plates are unsculptured and others have small coni. The specimen identified as Deflandrea dartmooria (P 25948) f r o m the Rivernook bed of the Dilwyn Clay (Cookson & Eisenack, 1967) lacks the linear markings at plate boundaries as well as indications of tabulation in the sulcus. In other respects, it shows the characteristics of D. medcalfii and is here transferred to that species. Types: Holotype, specimen illustrated by Cookson & Eisenack (1965, Plate 16, Figs 1, 2). No paratypes. Type locality: North bank of Glenelg River, southeast corner of the Parish of Drajurk, 14.5 km southeast of Casterton, Victoria. Type stratum: Carbonaceous mudstone near the base of the Dartmoor Formation, Early Eocene Malvacipollis diversus Zone. Additional occurrence: Sidewall 1,431.0 m in Tuna-3 (Loc. 20).

core

at

Deflandrea truncata sp. nov. (Plate 5, Figs 2, 3a-c) Material: Twenty-three specimens. Description: Periphragm is elliptical

to

Fig. 2. Arrangement of sulcal plates (unshaded) in Deflandrea

dartmooria.

PALAEOCENE AND EOCEr 3 SPECIES OF DEFLANDREA ovoid in dorso-ventral view and has an obtusely rounded apical margin and a truncated, straight to slightly concave antapical margin. Surface 1-1.5/* thick; smooth, finely pitted, or granulate (sculpturing evenly distributed). Cingulum is poorly defined by a shallow transverse band about 6/* wide located just below midheight. Ventrally, the cingulum is not offset and is interrupted by an ill-defined sulcus. Cingulum and sulcus discernible on about half the specimens. Periphragm may be thickened slightly at the apex and a thickening or a nipple-like projection may occur at each antapical corner. Outline of the endoblast is circular or nearly so in dorso-ventral view; endophragm is smooth and as thick as or slightly thicker than the periphragm. Endoblast occupies a considerable part of the pericoel. Intercalary archeopyle trapezoidal and the opening in the endophragm is nearly as large as that in the periphragm but usually appears shorter due to foreshortening. Operculum is free and isolated. Opercula usually consist of adherent pieces from both wall layers. Archeopyle is 34-40/* wide and 16-22/* long (width consistently about twice the length). Entire specimens are 86-104/* long and 6886/* wide (mean length :width ratio is 1:0.8). The nearly circular endoblast is 65-78/* long and 66-79/* wide. Usually the length-width dimensions on any specimen will be within 5/* of each other. Measurements taken on 10 specimens. Comparison: Deflandrea truncata differs from D. leptodermata by having less well developed horns, a straight or slightly concave antapical margin and a more elongate outline in dorso-ventral view. Types: Holotype (Plate 5, Figs 3a-c); Paratype (Plate 5, Fig. 1); another paratype not illustrated. Type locality: Locality 16. Type stratum: Flounder Formation at 2,198.1 m, Early Eocene Malvacipollis diversus Zone. Additional occurrences: Flounder-3 well (Loc. 16), at 2,189.9-2,199.1 m; Flounder-2 (Loc. 15) at 2,126.8 m. Deflandrea leptodermata Cookson & Eisenack, 1965 (Plate 5, Figs la, lb) 1965 Deflandrea leptodermata Cookson & Eisenack, pp. 121-122, Plate 11, Figures 6, 7 [1965a].

111

Material: Holotype, paratype, and 11 unillustrated specimens from the type locality. Comments: Periphragm and endophragm are smooth and the only indications of tabulation are the archeopyle, the cingulum and on some specimens, a faintly delimited sulcus. The cingulum is a shallow depression, offset ventrally, and the ends of the cingulum tend to curve antapically and merge with the margins of the sulcus. On some specimens (Plate 5, Fig. lb) a very low transverse ridge occurs along the middle of the cingulum. Archeopyle is exceptionally broad and there is little difference in the lengths of pieces of the periphragm and endophragm which form the operculum. The granulate sculpture on the cyst walls, stated in the original description of the species, resulted from the overoxidized condition of the type specimens. Types: Holotype, specimen illustrated in Cookson & Eisenack (1965A, Plate 11, Fig. 6). Paratype, specimen illustrated in the same publication (Plate 11, Fig. 5). Type stratum: Turritella beds of the Browns Creek Clays, 4.6 m below the Notostrea greensand: Middle to Late Eocene, Nothofagidites asperus Zone. Deflandrea phosphoritica Eisenack, 1938 (Plate 4, Figs 3a, 3b, 4a, 4b) 1965 Deflandrea phosphoritica Eisenack, Cookson & Eisenack, p. 121, Plate 11, Figure 11. 1965 Deflandrea granulosa Cookson & Eisenack, Plate 11, Figures 8, 9. Material: Between 75 and 100 specimens plus the holotype and paratype of Deflandrea granulosa. Comments: The brief synonymy above lists only the reported occurrences of the species in southeastern Australia. Deflandrea phosphoritica has also been recorded from Western Australia (Cookson & Eisenack, 1961) and from many localities outside Australia. Lengthy synonymies are given in Gocht (1969) and in Davey et al. (1966). On well-preserved specimens, the endophragm is thicker than the periphragm and has an even, finely granulate or faintly pitted sculpture. In contrast, the periphragm is smooth except for clusters of minute coni which correspond to the positions of plates. Based on the Australian specimens, the presence of 7 precingular, 5 postcingular and

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LEWIS E. STOVER

178

2 antapical plates can be demonstrated. Other plates that can be recognized include 2a, 1' and 4'. The cingulum is undivided. Outline of the periphragm in dorso-ventral view is perhaps the most variable feature among specimens of D. phosphoritica, with the outline differing from decidedly five-sided to roundly triangular with a slight antapical concavity. The prominence of the horns is least among the roundly triangular forms.

archeopyle or operculum are usually recognizable on the periphragm and other indications of tabulation are vague or lacking. In outline, the periphragm is typically tricornate, as in the specimen shown on Plate 4, Figures 5a-c, whereas on some specimens the horns are less pointed, which has the effect of giving the cysts a more rounded outline somewhat like that in specimens of Defiandrea leptodermata.

Additional occurrences: Defiandrea phosphoritica is the most widespread and one of the most commonly occurring species of Defiandrea in southeastern Australia. In the Browns Creek Clays (Loc. 7) the species occurs throughout the 30.8 m section from the base of the Turritella beds to the top of the outcrop. Other occurrences are: Perch-1 (Loc. 19) at 1,162 m, Dolphin-1 (Loc. 13) at 1,204.8-1,218.9 m, Turrum-1 (Loc. 21) at 1,979.5-2,006.9 m, and Groper-1 (Loc. 17) at 937.3-957.7 m in the Gippsland Basin; Bass-1 (Loc. 9) at 1,641.6 m in the Bass Basin. So far, D. phosphoritica has been found in Australia only in strata younger than Early Eocene.

Types: Holotype, specimen BX36 in the collection of Prof. G. Deflandre. Paratype, specimen illustrated by Deflandre & Cookson (1955, Plate 5, Fig. 6), endoblast only. Photomicrographs of the holotype are shown in the edition of Deflandre & Cookson (1955) re-issued by Deflandre et al. (1970). Type locality: Locality 8. Type stratum: Dark carbonaceous clay at the base of the Johanna River Sands immediately above the contact with the Cretaceous Otway Group: Middle to Late Eocene, Nothofagidites asperus Zone. Additional occurrences: Outwash exposure of the Browns Creek Clays (Loc. 7). The species occurs in samples between the base of the outcrop and the base of the Notostrea greensand (section given in Cookson & Eisenack, 1965a). Turrum-1 well (Loc. 21) at 1,954.7-2,031.3 m.

Defiandrea heterophlycta Deflandre & Cookson, 1955 (Plate 4, Figs 5a-c) 1955 Defiandrea heterophlycta Deflandre & Cookson, pp. 249-250, Plate 5, Figure 6, Text-Figure 5. 1965 Defiandrea heterophlycta Deflandre & Cookson, Cookson & Eisenack, Plate 12, Figures 1-4 [1965a]. 1970 Defiandrea heterophlycta Deflandre & Cookson, Deflandre et al, Plate 5, Figure 6, Text-Figure 5, also supplementary Plate 1, Figures 1-4. Material: Paratype; 7 specimens from the type locality and from the Browns Creek Clays (Loc. 7 ) ; and 23 specimens from Turrum-1 well (Loc. 21), Gippsland Basin, Victoria. Comments: The species is characterized by having a smooth, clear periphragm and a distinctly sculptured endophragm. Sculpture on the endophragm consists of irregularly shaped coarse grana and tubercules which vary in height from 1.5-7/x (most are 3/x or less) and in diameter from 2-5^. At high magnification the endophragmal surface between the grana and tubercules appears finely and uniformly pitted. The cingulum, sulcus and

Defiandrea extensa sp. nov. (Plate 5, Figs 4a-c, 5a-d, 6) Material: Between 200 and 300 specimens from several onshore and offshore Gippsland Basin wells. Description: Outline of periphragm in dorso-ventral view is broadly elliptical to ovoid with evenly convex lateral margins. Apical margin more convex than lateral margins and has a short apical horn at its apex; antapical margin concave with small horns developed at each corner (the right antapical horn is the larger). Periphragm is less than 1/x thick, spinose; spines mostly pointed, occasionally with bifurcate tips, 3-9/x long and fairly evenly spaced, except in the cingulum and sulcus which lack spines or have a few scattered spines. Endoblast in dorso-ventral view almost completely fills the pericoel. Endophragm, which is about 0.5/x thick and is close to the periphragm, has an outline conformable with that of the latter, deviating only slightly at the bases of the apical and antapical horns.

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA 179 Archeopyle intercalary, large, wider than taining the types also has an additional 160 long, has a trapezoidal outline with the base specimens of D. extensa. being wider than the apical margin. OperType locality: Locality 22. culum free. Tabulation is commonly obscure Type stratum: Latrobe Group at 85.4 m, although outlines of some precingular and postcingular plates can be identified by the Middle to Late Eocene, Nothofagidites asperus alignment of the spines, particularly on the Zone. dorsal surface of the epitract. Additional occurrences: Barracouta-3 (Loc. Specimens are 66-81^ long (mean 74/x) 12) at 1,099.2 m; Perch-1 (Loc. 19) at and 59-66^ wide (mean 62/x). Holotype is 1,143.8-1,161.4 m; Groper-1 (Loc. 17) at 72^ long, 65/x wide. Measurement based on 953.7-962.9 m. 25 specimens. Comparison: Deflandrea extensa differs from Deflandrea sp. cf. D. diebelii Alberti, 1959 D. longispinata Wilson, 1968 by being smaller (not illustrated) and by having shorter spines and shorter apiComments'. A few rather poorly preserved cal and antapical protrusions. Wilson described specimens were recovered (from Albacore-1 the archeopyle of D. longispinata as a four- (Loc. 11) at 2,795.0 which have strongly sided nearly equidimensional 2a plate. In D. attenuated apical andm),antapical horns and extensa, the archeopyle and operculum (Plate STYLE B-l archeopyles*. The periphragm 2a) have narrower apical than antapical mar- a somewhat striated appearance owing has gins and on some specimens, the archeopyle numerous longitudinal folds; the periphragmto tends to be six-sided. and endophragm are each less than thick. Types: Holotype (Plate 5, Figs 4a-c), Except for the archeopyle and a vaguely deparatypes (Plate 5, Figs 5a-d, 6). Slide con- fined cingulum, tabulation is absent. Too few SHAPE VARIATIONS IN TERTIARY SPECIES OF DEFLANDREA

Fig. 3. Variability in periblast outline. A, Deflandrea truncata; B, D. druggii; C, D. conorata; D, D. dilwynensis; E, D. pachyceros; F, D. flounderensis; G, D. medcalfii; H, D. pentaradiata. * See Fig. 5 and later text. Spec.Publs geol^oc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

LEWIS E. STOVER COMPLETENESS OF TABULATION

1/ F G M H Fig. 4. Completeness of tabulation. A, Defiandrea bakerii; B, D. pentaradiata; C, D. pachyceros; D, D. delineata; E, D. flounderensis; F and G, D. dartmooria; H, D. phosphoritica. Drawings not to scale; tabulation shown by unshaded areas and only the tabulation on the dorsal surface is depicted, except for G, which shows the tabulation on the ventral surface. specimens were recovered to evaluate varia- portrays most of the variability. In shape, bility and some morphologic features were periblasts vary from broadly or narrowly obscured by the dark coloration of the tracts. rounded, nearly elliptical with little or no Associated spores and pollen indicate the horn development to somewhat more elongate specimens are from the middle Palaeocene and tricornate with conspicuous horn development. Examples of the former are shown in Lygistepollenites balmei Zone. Figures 3A, B and those of the latter in Figures 3E, H. Species whose periblast outline is inDISCUSSION termediate between the two extremes are Variability shown in Figures 3C, D. In general, the Some appreciation of the expression and more rotund species tend to lack prominent degree of variability shown among Early horns and when the horn development is weak, Tertiary species of Defiandrea from south- the apical horn tends to be more clearly deeastern Australia may be gained by examining fined than the antapical horns. The completeness of tabulation on the periseparately the following features: 1, outline of the periblast in dorso-ventral view and the phragm remains fairly constant for a species, prominence of the apical and antapical horns; with one or two possible exceptions, but varies 2, the completeness of tabulation; 3, the widely among species. As illustrated by Figure archeopyle; 4, the position and size of the 4, the only indications of tabulation on some endoblast in relation to the periblast; and 5, species are the archeopyle or the cingulum (see Figs 4A, B, respectively). On other species, types and distribution of sculpture. Outlines of the periblasts for eight species the archeopyle, cingulum, and some of the of Defiandrea are depicted in Figure 3, which precingular plates are revealed as shown in

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA

Figures 4C, D. Still other species show more of the tabulation with some of the postcingular and some of the apical plates distinguishable, (Figs 4E, H), until the ultimate is reached in which the complete tabulation is discernible (see Figs 4F, G). On Deflandrea dartmooria, even some of the sulcal plates can be seen. Although the sizes and shapes of the plates are in some cases preserved with remarkable fidelity, rarely is the tabulation expressed conspicuously, and it can usually be resolved with certainty only under magnifications of at least 800. Among the Early Tertiary species, from the coastal and offshore basins of Victoria, Deflandrea phosphoritica is the only species thus far encountered in which the completeness of tabulation varies considerably, even among excellently preserved specimens. Although many specimens of D. phosphoritica have groups of minute coni denoting the positions of 1', 1", 3", 4", 5", 7", 1'", 3"', 5"', as well as the cingulum and operculum, it is not unusual to find specimens with smooth periphragms in which only the cingulum and oper-

ARCHEOPYLE STYLES STYLE A

181

culum or archeopyle are evident. On the other hand, relatively few specimens have been recovered in which nearly all of the tabulation, except for the anterior intercalary plates la and 3 a, is revealed. In general however, the majority of specimens show little of the tabulation. All species of Deflandrea possess intercalary archeopyles (Type I of Evitt, 1967), formed by the release or opening of plate 2a. For the purpose of this discussion, two styles of Type I archeopyle are recognized and termed informally STYLE A and STYLE B. STYLE A is characterized by its small size relative to the width of the periblast and is commonly elongate. Most commonly, the operculum is hinged at its base and therefore, remains attached (Figs 5B, C). Species with free or detached opercula (Fig. 5A) are much less common. Among forms with the STYLE A archeopyle, corresponding openings in the endophragm are rare. In contrast, STYLE B archeopyles are large, broad, do not remain attached, and there is always a corresponding opening in the endophragm. Almost STYLE B STYLE B-1

STYLE B - 2

Fig. 5. Archeopyle styles. A, operculum free, no opening in endophragm; B, operculum attached, no opening in endophragm; C, operculum attached, opening in endophragm; D. operculum free, convexly triangular; E, operculum free, hexagonal. Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

182

LEWIS E. STOVER PERIBLAST/ENDOBLAST

RELATIONSHIP

Fig. 6. Periblast/endoblast relationships. A, Deflandrea bakerii; B, D. delineata; C, D. medcalfii; D, D. pachyceros; E, D. pentaradiata; F, D. dilwynensis; G, D. heterophlycta; H, D. leptodermata; I, D. truncata.

without exception, the two pieces of wall periphragm, departing from the latter only at material of the operculum remain together. or near the bases of the apical and/or antaShape of STYLE B archeopyles is roundly pical horns, as depicted in Figure 6C and triangular (Fig. 5D), or irregularly hexagonal particularly Figures 6D, E. The position, size, with the width greater than the height (Fig. and shape of the endophragm itself, as well 5E); the former is referred to as STYLE B-l, as its relation to the periphragm, seems to the latter as STYLE B-2. Usually, the shape vary slightly within a species. The greatest and proportion of the archeopyle are constant variability appears to be among species in for a species, although the size may vary some- which the apical and antapical margins of the what depending on the overall size range of endophragm tend to protrude into the horns, the species. Reliable measurements of the e.g., in Deflandrea obliquipes. In such species archeopyle may be difficult to obtain unless the outline of the endophragm varies accorda large number of favourably oriented speci- ing to the amount of protrusion. Sculpturing is subdued on most species; mens are available. In dorso-ventral view, the endoblast gener- rarely is it conspicuous. If coarse or fairly ally occupies a position about midway between coarse, a species has usually a single type of the anterior and posterior margins of the sculpture; whereas, if fine, several types may cyst. The endoblast may fill a relatively small occur within a species; in such species indipart of the pericoel as shown in Figures 6A, B, vidual specimens almost invariably have just or a large part, as shown in Figures 6F-I. In a single sculptural pattern. Although an occaa majority of species, the outline of the endo- sional species has a sculptured endophragm, phragm shows little relationship or conformity sculpture on the periphragm is unequivocally to that of the periphragm; however, on a few more common, and in the latter it may be species the outline of a major part of the uniform or unevenly distributed. Because the endophragm follows quite closely that of the same sculptural pattern occurs on several

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA 183 species, sculpturing per se is of little taxo- weak, moderate, or strong and recording the nomic value. stratigraphic occurrences of the species in each category, the plot shown in Figure 8 STRATIGRAPHIC IMPLICATIONS emerges. Forms with strong horn developBy considering the various characteristics of ment are concentrated in the Palaeocene and Deflandrea stratigraphically, certain features Early Eocene; concomitant with the reduction tend to be more prevalent among species from in horn development is the tendency for the a particular part of the Early Tertiary section. inner body to occupy a larger portion of the As indicated in Figure 7, species having STYLE cyst. This tendency is depicted by the drawings A archeopyles occur only in Palaeocene. In on the right side of Figure 8 which show contrast, species having STYLE B archeopyles the outlines of the periphragms and endooccur in Palaeocene and Eocene strata. A fur- phragms of four species. Among Middle and ther differentiation is seen in that species with Late Eocene species the endoblast almost comSTYLE B-l archeopyles do not occur in strata pletely fills the pericoel and there is an inyounger than Early Eocene. Thus, all Middle clination for the periphragm outline to be to Late Eocene species of Deflandrea from south- rounded. It is also customary for these species eastern Australia have STYLE B-2 archeo- to have generally poorly delimited cingula and pyles. Also important is the fact that approxi- sulci; however, these are features more akin to mately three-quarters of the species, as shown tabulation. by the numbers in the bars of Figure 7, have The distribution of the species of Deflandrea STYLE B archeopyles. The following com- having STYLE B archeopyles is arranged ments refer just to those species with this according to the completeness of tabulation as archeopyle style. shown in Figure 9. Although no clear-cut By categorizing the horn development as picture emerges, there seems to be a tendency

Fig. 7. Stratigraphic distribution of archeopyle styles. Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

LEWIS E. STOVER

184

STYLE B ARCHEOPYLE

- DISTRIBUTION AND HORN DEVELOPMENT re£

HORN DEVELOPMENT

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PALEOCENE

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EARLY EOCENE

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Fig. 8. Stratigraphic distribution of species with STYLE B archeopyles arranged according to the prominence of the apical and antapical horns. for species with complete or nearly complete tabulation to occur in the Eocene. D. medcalfii is the only Palaeocene species possessing nearly complete tabulation but this form occurs also in the Early Eocene. In Palaeocene and Early Eocene species, specimens of a taxon are fairly constant as to how much of the tabulation they show, whereas among Middle and Late Eocene specimens of certain species, especially D. phosphoritica and to a lesser extent D. extensa, the amount of tabulation revealed by individual specimens varies and appears to be independent of preservation. Little relationship is seen between the type, uniformity, or intensity of sculpturing—or the lack of it—on either the periphragm or endophragm and the stratigraphic distribution of species, with possibly two exceptions. One is that species in which the periphragm is unsculptured or that have somewhat more intense sculpture on either the periphragm or endophragm have been identified from only

Middle to Late Eocene sections. Secondly, species with more or less uniform sculpturing, regardless of archeopyle style, occur consistently in the Palaeocene and less commonly in the Early Eocene. DISTRIBUTION AND COMPLETENESS OF TABULATION LATE AND MIDDLE EOCENE

s

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1

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Fig. 9. Stratigraphic distribution of species witb STYLE B archeopyles arranged according to the completeness of tabulation.

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA SUMMARY AND CONCLUSIONS Among Palaeocene and Eocene species of Deflandrea morphologic characteristics are generally fairly constant for a species, but differ extensively between species. In brief, the outline of the periblast in dorso-ventral view varies from narrowly to broadly rounded to somewhat triangular with attenuated corners. In general, the greater the triangularity, the better the horns are delimited. Tabulation varies from being excellently indicated, in which case all or nearly all of the plates are clearly shown, to being poorly indicated, in which case only the archeopyle or the archeopyle and cingulum are shown. Basically, two styles of archeopyle Type I are recognized: STYLE A is small, commonly elongate and its operculum is usually hinged; STYLE B is large, broad, has a corresponding opening in the inner body, and its operculum is usually free. The outlines of the endoblast and periblast vary from dissimilar to fairly conformable and the endoblast may occupy only a small part of the periblast or it may fill it nearly completely. Finally, sculpturing, which is generally inconspicuous, may cover the entire surface or it may be confined to specific areas, and it may occur on either body, but is usually on the periblast. Species of Deflandrea with STYLE A archeopyles occur only in the Palaeocene, whereas those with STYLE B archeopyles occur throughout the Early Tertiary. However, among the latter, species with the STYLE B-2 archeopyle are found only in assemblages from the Middle and Late Eocene. It seems, then, that with decreasing geological age, the height of STYLE B archeopyles tends to decrease and, concomitantly, the shape of the periblast becomes more rotund. Species with well-developed apical and antapical horns occur mainly in the Palaeocene and Early Eocene, and the prominence of the horns decreases with decreasing geological age. With the reduction in horn development, the endo-

185

blast tends to fill a greater amount of the periblast, so that in at least one Middle to Late Eocene species, there is practically no separation between the two bodies of the cyst. Tabulation is expressed best among the Early Eocene species and is generally vaguely and poorly reflected on Palaeocene and Middle to Late Eocene forms. Finally, intense sculpturing has been observed only on Middle-Late Eocene species. Knowledge of these morphological features —their variability, modes of expression, prevalence of occurrence, and relationships to each other—in conjunction with reliable stratigraphic data should lead eventually to determining phylogenetic lineages for species of Deflandrea. But until that time comes, information now available can be used advantageously in defining and assessing new taxa and utilizing all species for dating sections in which other palaeontological data are lacking or inconclusive. ACKNOWLEDGMENTS Appreciation is expressed to Esso Australia Ltd and Broken Hill Proprietary Co. Ltd, for permitting the release of the data derived from offshore Gippsland and Bass Basin wells and for contributing to the cost of publication. I am also most grateful to T. A. Darragh and K. N. Bell of the National Museum of Victoria, Melbourne, for making available the type specimens of Deflandrea in their collection and for the courtesies extended to me while in Melbourne. I also wish to thank D. J. Taylor of the University of Sydney, who accompanied me in the field and contributed provocative and stimulating discussions on the Tertiary stratigraphy of southwestern Victoria. Last but not least, I am most appreciative to my colleagues at the Esso Palynology Laboratory—Dr P. R. Evans, A. D. Partridge, and Mrs A. L. Nicholls—for bringing numerous specimens to my attention and aiding in the preparation of the manuscript.

REFERENCES I. C . , 1956: Additional microplankton from Australian Late Mesozoic and Tertiary sediments. Aust. J. Mar. Freshwat. Res., 7, pp. 183-191. , & EISENACK, A., 1958: Microplankton from Australian and New Guinea Upper Mesozoic sediments. Proc. R. Soc. Vict., 70, pp. 19-79.

COOKSON,

Spec.Publs geol.Soc.Aust, 4: pp. 167-188, Pis 1-5, 1973.

—, , 1961: Tertiary microplankton from the Rottnest Island bore, Western Australia. J. Proc. R. Soc. West. Aust., 41, pp. 39-47. —, , 1965a: Microplankton from the Browns Creek Clays, SW. Victoria. Proc. R. Soc. Vict., 79, pp. 119-131.

LEWIS E. STOVER , , 19656: Microplankton from the EISENACK, A., 1938: Die Phosphoritknollen der Bernstein-formation als Uberlieferer TerDartmoor Formation, SW. Victoria. Proc. R. tiares Plankton. Schr. phys.-dkon, Ges. Soc. Vict., 79, pp. 133-137. Konigsbg., 70 (2), pp. 181-188. , , 1965c: Microplankton from the Paleocene Pebble Point Formation, south- EVITT, W. R., 1967: Dinoflagellate studies II. The western Victoria, Pt 2. Proc. R. Soc. Vict., 79, archeopyle. Stanford Univ. Pubis, Geol. Sci., pp. 139-146. 10 (3), pp. 1-82. , , 1967: Some microplankton from the Paleocene Rivernook Bed. Proc. R. Soc. GOCHT, H., 1969: Formengemeinschaften altterVict., 80, pp. 247-257. tiaren Mikroplankton aus Bohrproben des DAVEY, R . J., et al., 1966: Studies on Mesozoic Erdolfeldes Meckelfeld bei Hamburg. and Cainozoic dinoflagellate cysts. Bull. Br. Palaeontographica, 126B, pp. 1-100. Mus. Nat. Hist., Geol, suppl. 3, pp. 1-248. DEFLANDRE, G . , & COOKSON, I. C . , 1 9 5 5 : Fossil HARRIS, W. K., 1965: Basal Tertiary microfloras microplankton from Australian Late Mesozoic from the Princetown Area, Victoria, Australia. and Tertiary sediments. Aust. J. Mar. Palaeontographica, 115B, pp. 76-106. Freshwat. Res., 6, pp. 2 4 2 - 3 1 3 . DEFLANDRE, G . , et al., 1970: Re-issue of Deflandre SINGLETON, O. P., 1967: Otway Region; in & COOKSON, 1955 (in French), by LaboraMcAndrew, J., & Marsden, M. A. H. (Eds), toire de Micropaleontologie de l'ficole PracGeology Excursions Handbook, pp. 117-131. tique des Hautes Etudes, Institut de PaleonA.N.Z.A.A.S. 39th Congress, Melbourne. tologie du Museum, Paris; avec addendum et WILSON, G. J., 1967: Microplankton from the postface, pp. 1-70, 1-54. Garden Cove Formation, Campbell Island. DOUGLAS, J. G., 1960: Microplankton of the DeN.Z.JI Bot., 5, pp. 223-240. flandreidae Group in western district sediments. Min. geol. J., 6 (4), pp. 17-32. , 1968: Palynology of some Lower TerDRUGG, W. S., 1967: Palynology of the Upper tiary Coal Measures in Waihao District, Moreno Formation (Late CretaceousSouth Canterbury, New Zealand. N.Z.JI Bot., Paleocene), Escarpado Canyon, California. 6, pp. 56-62. Palaeontographica, 120B, pp. 1-71. *Present Address: Lewis E. Stover* Production Research Co., Esso Australia Ltd, P.O. Box 218, G.P.O. Box 4047, Houston, Texas 77001, Sydney, U.S.A. New South Wales 2001.

186

PALAEOCENE AND EOCENE SPECIES OF DEFLANDREA APPENDIX TYPES Specimens illustrated on Plates 1-5 or designated as types are listed below. These specimens are in the palaeontological collections of the National Museum of Victoria, Melbourne. Slides have been assigned accession 'P' numbers and the exact locations of the specimens are on file with the collection. 'P' Number Figure Name Plate Type Loc. P28100 D. druggii 3 Holotype 15* 1 P28122 2 D. bakerii Hypotype 1 D. conorata P28123 8 1 Holotype 16 P28124 D. dilwynensis 5 4 Hypotype 1 D. dilwynensis P28125 1 6 4 Hypotype D. medcalfii 4 Not illustrated Paratype D. dartmooria P28126 20 4 2 Hypotype D. extensa P28127 5 4 Holotype 22 D. extensa 22 Paratype 5 5 D. extensa 22 Paratype 5 6 P28128 D. flounderensis Holotype 14 3 1 P28129 D. flounderensis 2 Paratype 16 3 P28130 D. heterophlycta 4 8 5 Hypotype P28131 D. leptodermata 7 Hypotype 5 1 P28132 D. medcalfii 4 3 Holotype 3 P28133 D. obliquipes 2 7 Hypotype 1 D. pachyceros 2 2 Hypotype 1 P28134 D. obliquipes 1 2 Hypotype 8 D. pachyceros Hypotype 2 1 1 P28135 D. pachyceros Hypotype 1 2 3 P28136 D. phosphoritica 7 4 Hypotype 3 P28137 D. phosphoritica Hypotype 7 4 4 P28138 D. truncata Holotype 16 5 3 P28139 D. truncata Paratype 15 5 2 P28140 D. truncata Paratype 16 Not illustrated D. druggii P29902 Paratype 16 1 4 Previously designated type specimens D. pellucida P16237 Holotype 1 7 D. robusta PI6242 Holotype 2 5 D. bakerii P23081 Hypotype 1 1 P24068 D. pentaradiata Holotype 2 6 D. delineata P24071 Paratype 2 4 P24080 D. dartmooria Holotype 3 4 D. dartmooria Holotype 4 1 * Specimen on slide provided to the National Museum of Victoria by Dr I. C. Cookson and labelled 'Pebble Pt D. bakerii'. — — —

—

—

—

—

EXPLANATION OF PLATES PLATE 1

Figs 1, 2. Deflandrea bakerii Deflandre & Cookson, 1955. 1, hypotype shown in bright field and focused on the dorsal surface (la) and about midway through the specimen (lb), X400. 2, Hyptotype in interference contrast showing sculptural details, operculum outline and smooth endophragm, XI000. Figs 3, 4. Deflandrea druggii sp. nov. 3, Holotype shown in interference contrast (3a) and bright field (3b). X400. 4, Paratype in bright field, X400. Figs 5,6. Deflandrea dilwynensis Cookson & Eisenack, 1965. 5, Hypotype shown in bright field, XI000. 6, Another hypotype shown in bright field (6a) and phase contrast (6b); the latter shows the operculum outline, X625. Specimens are practically colourless, thin-walled and uncompressed. Fig. 7. Deflandrea pellucida Cookson & Eisenack, 1958. Holotype; antapical part in bright field (7a) and antapical part of specimen in interference contrast showing grana, archeopyle and endophragm (7b), X500. Fig. 8. Deflandrea conorata sp. nov. Holotype shown in bright field (8a) and interference contrast (8b) at the same focus level, X400. Spec.Publs geol.Soc.Aust., 4: pp. 167-188, Pis 1-5, 1973.

LEWIS E. STOVER PLATE 2

Figs 1-3. Deflandrea pachyceros Deflandre & Cookson, 1955. 1, Hypotype showing openings in both the periphragm and endophragm (la) and the rather even sculpturing on the ventral surface (lb), bright field, X625. 2, Detail of verrucae on another hypotype, XI000. 3, Another hypotype shown in interference contrast and focused on the dorsal (3a) and ventral (3b) surfaces, X625. Fig. 4. Deflandrea delineata Cookson & Eisenack, 1965. Mid-dorsal part of the paratype showing the operculum and surface of the periphragm in interference contrast, X400. Fig. 5. Deflandrea robusta Deflandre & Cookson, 1965. Holotype in interference contrast, X400. Fig. 6. Deflandrea pentaradiata Cookson & Eisenack, 1965. Holotype, X400. Figs 7,8. Deflandrea obliquipes Deflandre & Cookson, 1955. 7, Sculpturing on the mid-ventral surface of the periphragm of a hypotype. 8, Bright field photomicrographs of another hypotype focused on the ventral surface (8a) and about midway through the specimen (8b), X625. Figure 8c shows the sculpturing on the ventral surface in interference contrast, XI000. PLATE 3

Figs 1,2. Deflandrea flounderensis sp. nov. 1, Holotype shown in consecutive focus levels in bright field from the ventral (la) to the dorsal surface (lc), X500. 2, Paratype shown in bright field (2a) and interference contrast (2b) at the same .focus level, X500. Fig. 3. Deflandrea medcalfii sp. nov. Holotype shown in interference contrast focused at about midway through the specimen (3a) and on the dorsal surface (3b). Figure 3 c shows the dorsal surface of the hypotract and part of the epitract, X500. Fig. 4. Deflandrea dartmooria Cookson & Eisenack, 1965. 4a, 4b, Slightly different focus levels of the ventral surface showing part of the tabulation and the sulcal scar in interference contrast, X500. PLATE 4

Figs 1,2. Deflandrea dartmooria Cookson & Eisenack, 1965. 1, Various focus levels showing parts of the holotype: la, dorsal surface of apical horn and archeopyle; lb, endophragm and periphragm at about midway through the specimen; lc, Id, views of mid-dorsal surface showing antapical margin of the archeopyle, cingulum, and tabulation, X400. 2, Ventral surface of paratype showing plates 1', 1", 7", V" and 5'" as well as some of the sulcal plates and the sulcal scar, X400. Figs 3,4.Deflandrea phosphoritica Eisenack, 1938. 3, 4, Paratypes showing some of the variability in shape in bright field, X500. Fig. 5. Deflandrea heterophlycta Deflandre & Cookson, 1955. Topotype in bright field and focused on the dorsal surface (5a), about midway through the specimen (5b), and on the ventral surface (5c), X500. Fig. 1.

PLATE 5

Deflandrea leptodermata Cookson & Eisenack, 1965. Hypotype shown in bright field (la) and in interference contrast (lb), X400. Figs 2, 3. Deflandrea truncata sp. nov. 2, Paratype shown in bright field (3a) and in interference contrast showing the dorsal (3b) and ventral surfaces (3c), X500. Note operculum inside holotype. Figs 4-6. Deflandrea extensa sp. nov. 4, Holotype, part of the dorsal surface (4a), at about midway through the specimen (4b), and most of the ventral surface (4c). 5, Paratype, specimen with shorter spines than the holotype and weaker horn development; dorsal surface (5a, b), about midway through the specimen (5c) and the ventral surface (5d). 6, Paratype, outline of hypotract about midway through the specimen showing antapical horns. Specimens shown in interference contrast, all X625.

LEWIS

E.

STOVER

Spec.Pubis geol.Soc.Aust., 4, 1973.

PLATE 1

PLATE

2

LEWIS

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STOVER

LEWIS

E.

STOVER

Spec.Pubis geol.Soc.Aust., 4, 1973.

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PLATE

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LEWIS

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5a Spec.Pubis geoI.Soc.Aust., 4, 1973.

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5

STATISTICAL APPROACHES TO POLLEN ANALYSIS

THE NUMERICAL ANALYSIS OF MODERN POLLEN SPECTRA FROM NORTHEAST QUEENSLAND RAIN-FORESTS By A. P. KERSHAW (With 2 Tables and 3 Text-Figures) ABSTRACT

Limitations to the interpretation of two pollen diagrams derived from volcanic crater lake deposits on the Atherton Tableland prompted this study of present-day pollen deposition within rain-forests. Pollen counts were obtained for present-day moss samples collected in fourteen rain-forest sites selected to cover a large range of the major environmental variables: altitude, annual rainfall and soil parent material. Numerical methods were employed to help analyse floristic and pollen data for the sites and to facilitate comparisons between them and with the environment. Comparisons were then made between different numerical treatments of the pollen data and one treatment was decided upon which, it is suggested, would be most useful in an objective comparison with fossil spectra from the pollen diagrams. INTRODUCTION Two pollen diagrams produced from volcanic crater lake deposits on the Atherton Tableland (Kershaw, 1970, 1971), in an area which under undisturbed conditons was covered by rain-forest, have been interpreted on the relative proportions of a few selected pollen taxa. Because of a limited knowledge of the pollen production and dispersal of the plants involved, these interpretations must be considered very tentative. An examination of modern pollen rain was therefore undertaken to enable better interpretation of the diagrams. Other studies in this field have tended to concentrate on a direct comparison between the composition of defined vegetation communities and pollen spectra derived from them and, in the most detailed analyses, 'R factors' have been produced which express the ratios of pollen percentages within the pollen sum to species percentages in the vegetation (Davis, 1963). This direct approach has proved very useful in areas with a fairly simple vegetation but there are several major problems for its adoption in a tropical rain-forest situation. These are stated below. 1. The vegetation tends to change continuously except where there is a sharp change in topography or soils. 2. The richness of the flora and absence of dominant individual species necessitate the identification of many pollen taxa and this would be virtually impossible at the species level. Spec.Publs geol.Soc.Aust., 4: pp. 191-199, 1973.

3. The complexity of the forest structure precludes easy identification of all plant species (particularly lianes and epiphytes growing in the canopy) that would be contributing to a pollen sum. It was therefore decided that some alternative method of comparison was desirable. Webb et al. (1967a) made use of numerical methods to examine floristic data from north Queensland rain-forest sites, and were able to correlate floristic groups with different gradients of inferred soil nutrient status, moisture and altitude, and to a large extent, with an intuitive classification of the vegetation using structural features (Webb, 1959, 1968). The present study seeks to determine the relationships of such floristic groups with the botanical composition of the present-day pollen fallout in a number of rain-forest plots. GEOGRAPHICAL SETTING Major features of the area selected are shown on Figure 1, which was compiled from 1:250,000 Military Survey topographical maps and Department of National Development geological maps, a rainfall map published by the Ministry of National Development in 1950, and an isohyet map supplied by the Queensland Irrigation and Water Supply Commission. The area contains the most accessible forest variety in tropical Queensland. Rain-forest covers most of the region but is replaced by open sclerophyllous vegetation under rainfalls of less than about 50-60 ins (ca 1,400 m m ) ,

A. P. KERSHAW 192 and on soils with impeded drainage or of low frequently covers the mountain tops during nutrient status. The transition from rain-forest part of the relatively dry days of winter and to open sclerophyll vegetation is normally very also influences the vegetation at higher altiabrupt except where there is a belt of tudes. 'wet sclerophyll forest' which characteristically has a canopy layer composed of sclerophyllous DATA species and an understorey of rain-forest species. Much true rain-forest originally cover- 1. The sites Fourteen sites covering a wide range of each ing the coastal areas has been cleared, mainly for sugar cane, while on the Atherton Table- of the three major environmental variables land the rain-forest has given way to dairy (altitude, parent material and annual rainfall) were selected. Their positions are shown on pasture and maize cultivation. The major physiographic features of the Figure 1 and briefly described in Table I: all study area trend approximately north-south are from rain-forest except sites 13 and 14 and are controlled largely by the underlying which are from wet sclerophyll forest. Sites geology (de Keyser, 1964). From the coast 4, 5, 6, 8, 9, 10, 11 and 12 are £ acre plots established by L. J. Webb and J. G. Tracey, inland they are: site 2 is an Atherton Forestry Office i acre 1. A discontinuous line of coastal ranges plot, 1, 3, 7 and 13 are L. J. Webb and composed of granitic and metamorphic J. G. sites Tracey spot lists and site 14 is a spot rocks varying in altitude from 150-1020 m. list compiled by J. Hopkinson. 2. The coastal plains which consist of river alluvia, swampy lagoonal deposits and old 2. Floristic data marine sediments. Floristic lists derived from plots are much 3. The central highland belt, composed mainly more complete than spot lists which name only of Palaeozoic metamorphics and intruded the canopy trees. In order to compare the two of list usefully only presence and absence Upper Palaeozoic granite. This contains the types data on the big tree species {i.e. those species highest peaks in Queensland, Mount Bartle capable reaching a trunk diameter of at least Frere (1608 m) and Bellenden Ker (1593 30 cm) ofwere used. The relevant species, which m). numbered were kindly listed by J. G. 4. The Atherton Tableland, which extends Tracey. The245,justification for reducing the from Mareeba to south of Millaa Millaa, floristic composition of the plots in this manner and formed from Late Cainozoic basaltic is found in Webb et al. (1967Z?) flows and associated pyroclastic deposits cluded from a comparative analysiswhoof conten (Best, 1960). It averages 800 m in altitude. species groups from the north Queensland 5. The Herberton Highland, largely composed rain-forest sites that 'the big tree species carry of granites and rhyolites rising to over the whole of the classificatory information'.

1200 m.

The soils supporting rain-forest are well 3. Pollen data Four moss sub-samples were collected from drained and derived from a variety of parent rocks. Generally, the complexity of the rain- rotting trees on the forest floor within each of the 14 sites. As moss was rare in most of forests decreases with decreasing nutrient stathe sites it was impossible to introduce a strict tus of the soils (Webb et al. 1970). sampling procedure. Sub-samples were collected The area embraces a wide range of climatic conditions. Rainfall is high along the coast at least 10 m apart. They were prepared for pollen analysis by a standard acetolysis method, and on the Central Highlands belt but declines sharply inland. It is markedly seasonal and most the pollen mounted in silicone oil (A.K. 2000) and examined under a Carl Zeiss (Oberfalls in summer. The average daily mean temperature for Cairns at sea level is 24°C, while kochen) automatic photomicroscope. Counts were continued until a total of 50 grains of the at Atherton at an altitude of 750 m it is 20°C. Occasional frosts occur between June and 'common taxa' had been recorded.* Four subAugust on the uplands above 600 m. Cloud sample counts from each site were compared, Common taxa are defined as those dry land pollen taxa contributing more than an average of one per cent to the pollen sum from all fossil spectra in the pollen diagrams from the Atherton Tableland They are Agathis, Podocarpus, Cunoniaceae (dicolpate), Cunomaceae (tncolpate), Elaeocarf

5

pus, Macaranga, Mallotus, Trema, Urticaceae-Moraceae, cf. Euodia, Casuarina and the Myrtaceae.

NUMERICAL ANALYSIS OF MODERN POLLEN SPECTRA 193 and each gross deviation in the representation SITE ANALYSES of a taxon in one count from the other three 1. Numerical methods was assumed to be due to very local influences and corrected to a value of the mean of the An information statistic has been used as other three values, before the counts were com- the basis of all computer programme analyses bined. One hundred and forty pollen taxa that were run on the CSIRO Control Data were separated including some that were not 3600 computer at Canberra. For the qualitative identified. The taxa probably represent all (presence and absence) data the polythetic life-forms existing in rain-forest as well as in- agglomerative program CENTCLAS (Williams troduced species and plants from the sclero- et al., 1966) and its monothetic divisive counphyll vegetation. Pollen from only 7 of the terpart DIVINF (Lance & Williams, 1968) big tree species included in the floristic site have been used. Quantitative pollen data have lists could be identified with certainty. been converted and analysed in percentage by CENTPERC (Dale et al., 1971; Dale The pollen data are analysed in four dif- &form Walker, 1970) and in the form of multistates ferent forms, listed below. (8 states taxon) by the agglomerative 1. Qualitative; presence and absence of all programmeto each MULTBET (Lance & Williams, taxa. 1967) and its divisive counterpart MULTDIV. 2. Percentaged; all taxa as percentages of MULTBET and MULTDIV also handled the total pollen. mixed pollen data with each quantitative entry 3. Multistates; actual numerical values of divided into 5 states. each taxon classed into a number of states. Inter-site similarity coefficients produced by 4. Mixed; classed states for common taxa and the agglomerative classificatory programmes presence and absence for other taxa. were used as input to the ordination proTABLE I

Site descriptions

Site

Locality

Structural form

1 Tinaroo Range Range 3 Sluice Creek 4 Davies Creek 5 Danbulla 6 Windin 7 Downfall Creek 8 Kuranda 9 Maalan turn off 10 McNamee Creek 11 Mirriwinni 12 El Arish

Simple notophyll — microphyll vine forest Simple notophyll— mixed notophyll vine forest Mixed notophyll vine forest Mixed notophyll vine forest + Agathis Simple notophyll vine forest Complex mesophyll vine forest Complex notophyll vine forest Mixed mesophyll vine forest Complex mesophyll vine forest Complex mesophyll vine forest Complex mesophyll vine forest Mixed mesophyll evergreen vine forest 13 Boar Pocket Wet sclerophyll forest Rd. 14 Tinaroo Road Wet sclerophyll forest 2 Herberton

Spec.Publs geoI.Soc.Aust., 4: pp. 191-199, 1973.

Approx. average annual rainfall (mm) (ins) 1900 75 1650 65

Altitude (m) (ft) 1160 3800 1130 3700

2540 1780 1650 3300 1500 2000 2670 3500 4300 3170

100 70 65 130 60 80 105 140 170 125

1040 760 760 760 700 425 520 150 60 60

3400 2500 2500 2500 2300 1400 1700 500 200 200

1500 1400

60 55

760 1070

2500 3500

Parent material granite granite basalt/ rhyolite granite metamorphics basalt granite metamorphics basalt basalt granite metamorphics metamorphics granite

194

A. P. KERSHAW 146° E

CAIRNS

ICURANDA^

QUEENSLAND CAIRNS BRISBANE 146° E ,MAREEB>

50RD0NVALEJ

l a k e ^ l£URAMOO

\

T1NAROO. . RESER VOIR

BELLENDEN KER

ATHERTON

QUINCAN CRATER '

LYNCHS CRATER

IERBERTON BROMFIELD SWAMP•

MTLLAA MILLAA

INNISFAIL

RAVENSHOE

•

11

Forest site Pollen analytical site Contours in feet

———

Isohyets in inches Quaternary sediments Atherton basalt Granites and rhyolites Metamorphics

jfc

TTT ^

EL A R I S H

5 Km

Fig. 1. Location of sites.

NUMERICAL ANALYSIS OF MODERN POLLEN SPECTRA DATA

PROGRAM

195

SITE GROUPS

FLORISTIC

CENTCLAS

7,13, 14

4,5,8,12

1,2,3

6,9,10,11

QUALITATIVE

DJVINF

7,12,13,14

4,5,8

1,2,3

6,9,10,11

1,2,4,7

3 , 6

10 ,11,12

1,2,4

3,6,9

10,11,12

2,4,5,7

1.3

6,9

10,11 ,12

2,4,5,8,13,14

7,11

1#3

5,7,8,13,14

1,2,3

13,14

4,5,7,8

1,2

3,6,9

10,11 ,12

5.7

1 2,3.4

6,9

10,11.12

POLLEN

CENTCLAS

1 3 , 14

5 ,8 , 9

QUALITATIVE

DIVINF

13 ,14

5,7,8

POLLEN PERCENTAGES

CENTPERC

8,13,14

POLLEN

MULTBET MULTDIV

MULTBET

MULTI STATES

POLLEN MIXED

MULTDIV

1 s t HIERARCHICAL DIVISION

10.12

10,11,12

2 n d HIERARCHICAL DIVISION

Fig. 2. Classifications on site data. Spec.Publs geol.Soc.Aust., 4: pp. 191-199, 1973.

6,9 4,6,9

A. P. KERSHAW 196 gramme GOWER using the method of Gower 2. Sites 6 and 9 which are both from basalt at medium altitude and have high rainfall. (1966). 3. The high altitude sites 1 and 2. 2. Classifications a. On floristic data. There is close corres- 4. The low rainfall sites 5 and 7. pondence between site groups formed from 5. The wet sclerophyll sites 13 and 14. both divisive and agglomerative classifications Site 3 can be attributed to group 2 or 3 terminated at the four group level (Fig. 2). in that it is at high altitude but on marginal The hierarchies produced are also very similar basalt. Site 4 is intermediate between groups with sites 6, 9, 10 and 11, all designated com- 3 and 4 in altitude and annual rainfall. Site plex mesophyll vine forest, separating from 8 is situated very near to a sclerophyll forest the less complex types at the highest level, then composed almost entirely of Casuarina littoa group containing the wet sclerophyll sites ralis. Where Casuarina pollen is treated in quan13 and 14 separating off with a final division titative form, except in the agglomerative between the high altitude sites, 1, 2 and 3 and classification on mixed data, it seems to have the remainder. Site 7 in both classifications an over-riding influence on the classification joins with sites 13 and 14 and this could be and the site is grouped with the 'wet sclerophyll because the spot list is small and possibly un- forests' rather than with the rain-forests under representative, and hence the site is speciespoor in comparison with others from the lower rainfalls. rain-forest. Site 12 is the only one that does not 3. Ordinations join the same group in both classifications. a. On floristic data. The first two Gower extracted are plotted graphically on b. On pollen data. There is little overall con- vectors Figure 3. The isolation of sites 6, 9, 10 and sistency in the hierarchical pattern produced 11 correlates with their separation at the from classifications on the different numerical first division well of the classification hierarchies treatments of the pollen data (Fig. 2). while the group formed of sites 7, 12, 13 and (i) At the highest level, division in both 14 supports the second division of the divisive classifications on the qualitative pollen data and classification. There is little of the the divisive classification of the mixed data formation of groups 1, 2, 3 suggestion and 4, 5, 8 but separates the lowland sites 10, 11 and 12 from these emerge in a plot of the 1st and 3rd the others. vectors. (ii) The classification of the sites on perOn pollen data. Again the first two veccentaged data and agglomerative classification torsb. are for each treatment of the of the sites on mixed data divide on annual pollen dataplotted (Fig. 3). The graphs for qualitarainfall. tive and mixed data produce very similar (iii) The classifications on the quantitative groupings (10, 11 and 12; 3, 6 and 9; 1, 2 and data separate the high rainfall and high 4; and 7, 8, 5, 13 and 14) which largely agree nutrient status forest sites from the others. with the overall groupings suggested by the Where sites 10, 11 and 12 are not separated classifications, and are almost exactly the same at the first division they divide off at the as the groupings produced by the divisive second, except that site 11 is missing from the classification of qualitative data. The results group in the agglomerative classification of the of ordination of the percentaged and quantiquantitative data. The only group not yet men- tative data do not give very meaningful site tioned that is formed at the second division is groupings, but in the graph of percentaged that composed of sites 13 and 14 in the ag- data the grouping of sites 6, 9, 10 and 11, glomerative classification of qualitative data. which is so consistent from the analyses of the All division has been stopped at the four floristic data, recurs. or five group level and here there is a good relationship between the groups formed by DISCUSSION the different classifications. Five site groups are 1. Floristic-pollen comparisons identifiable which can, to a certain extent, be The primary separation, from the analyses related to different measures of the major performed on floristic data, is of the high soil environmental variables. fertility and high rainfall sites 6, 9, 10 and 11, 1. The lowland high rainfall sites 10, 11 and while the remainder are subdivided into groups of high altitude, low general soil fertility and low 12.

NUMERICAL ANALYSIS OF MODERN POLLEN SPECTRA 1 10 • 11 I 3 I 10 • • 11 #

1

2

197 •

• 10

12

• 12

•

11 •

]2 13. *1

•9

# U

• 6

•4

• 2

/

4

•5

4

8•

I

Floristics

2

Pollen - qualitative

3

Pollen - percentages

4

Pollen - multistates

5

Pollen - mixed

*• 13

1 •

• 7

•10

7

13 • • 14

5

• 10

1

12 •

1st Vector

1

8

* 5 •

•3

•1

I

4 V

•6

3 V8 • 2

•

12 11 •

• 6 11 • •9 • 3

• 9

3

14 5 13* • 4 »8 • • • 7 2 • 1

•

6

2 • •

1

•

4 5 • 7 •

• 8

«14

13 •

Fig. 3. Ordinations on site data. rainfall. These groups are typically 'ecological' with edaphic factors having a primary influence. On the other hand, from analyses on pollen data, no marked edaphic control over group formation is evident. Pollen derived from a larger source area than a floristic plot, is likely to reflect vegetation from a variety of parent materials. In the case of the clear separation of the lowland sites, pollen contamination of sites 10 and 11 from the surrounding common metamorphics with vegetation of site 12 type may be important although the proximity of all three sites to edaphically controlled sclerophyll communities and cyclone damage may also have a significant influence. Sites 6 and 9, which combined with lowland sites 10 and 11 in the floristic classification, tend to remain together in the analysis of pollen data and not join any other group. Despite differences in individual site relationships between remaining sites, groupings produced on pollen data are, in general, not very different to those from floristic data. From the vectors produced by the Gower ordination it was possible to compare objectively the floristic pattern with those from the Spec.Publs geoI.Soc.Aust., 4: pp. 191-199, 1973.

various treatments of the pollen data. The Canberra program CANONGO (Williams & Lance, 1968) performs a simplified version of canonical analysis rotating the axes from two sets of vectors to get the best possible fit between them, and that is then expressed as a canonical correlation. The results obtained by comparing the first six floristic vectors with those from the pollen data are shown in Table II. All correlations are very good for at least the first three components, but the best correlation is with the mixed data which also show an 0.9 correlation between comparisons on the first four axes. 2. Comparisons between treatments of the pollen data It is important to know which treatment of the pollen data would best serve as a framework for comparison with fossil pollen spectra. The quantitative data obviously contain the greatest amount of information, but site groupings were less consistent between different kinds of analyses employed and not as easy to interpret as those produced from qualitative and mixed data. As fossil pollen spectra are derived from sediments that have accumulated

198

A. P. KERSHAW TABLE I I

Canonical correlations between vectors produced by ordination on floristic data and different treatments of the pollen data. 1i 1st Vector

2nd Vector

3rd Vector

4th Vector

5th Vector

6th Vector

Floristics/Pollen Qualitative

0-9867

0-9726

0-9246

0-7814

0-4365

0-1265

Floristics/Pollen Percentages

0-9914

0-9552

0-8021

0-7373

0-1866

0-0634

Floristics/Pollen Multistates

0-9900

0-9738

0-8574

0-6925

0-3165

0-0645

Floristics/Pollen Mixed

0-9959

0-9879

0-9711

0-9038

0-5644

0-2027

Site Correlations

outside the rain-forest canopy, they would tend to reflect a more regional vegetation picture than spectra derived f r o m material collected under the forest canopy, which contains many very local over-represented taxa. These taxa will be emphasised much more if treated as quantitative values. On the other hand, if all pollen data are in qualitative form, many important taxa, notably the common taxa, where changes in representation rather than presence or absence are important, will play little or no part in comparisons. It was found by an inverse analysis on the qualitative data that 7 out of the 12 common taxa were contributing no information to the site classification. The adoption of quantitative values for common taxa and qualitative values f o r other taxa, i.e. 'mixed' data as here used, may give more information f o r comparison with fossil spectra. CONCLUSIONS The classification and ordination of sites have produced groups which, though different f r o m floristic and pollen data, can be interpreted to a large extent in environmental terms. Site relationships f o r all treatments of the pollen data correlated well with those for floristic data

in canonical analysis. The treatment of pollen as mixed data exhibited the best correlation with the floristic data and would be most comparable with fossil pollen spectra. It should therefore be possible, if existing floristic associations have remained constant over the period of interest, to place fossil spectra within the framework produced by the modern spectra, and interpret changes in pollen diagrams f r o m within the area in fairly precise environmental terms. The absence or erratic representation of the pollen of many rain-forest plants need not inhibit the application of pollen analysis to rain-forest history.

ACKNOWLEDGMENTS I am very grateful to D r L. J. Webb, M r G. J. Tracey, D r J. Hopkinson, M r B. P. M. Hyland and M r E. Volck for information on vegetation sites and to D r M. B. Dale and M r P. Milne for valuable advice on the selection and running of computer programmes. Professor D. Walker, D r L. J. Webb and D r Suzanne L. Duigan read the manuscript and provided many useful comments, while M r M. Pancino drew the text-figures.

REFERENCES BEST, J. G., 1960: Some Cainozoic basaltic volcanoes in North Queensland. Rec. Bur. Miner. Resour. Geol. Geophys. Aust., 1960/78 [unpublished]. DALE, M .

B., LANCE, G . N . ,

& ALBRECHT,

1971: Extensions of information Aust. Comput. J., 3, pp. 29-34. DALE, M . B., & WALKER, D . , 1 9 7 0 :

L.,

analysis.

Information

analysis of pollen diagrams I. Pollen Spores, 12,

pp.

21-37.

DAVIS, M. B., 1963: On the theory of pollen analysis. Am. J. Sci., 261, pp. 897-912. DE KEYSER, F., 1964: Innisfail, Queensland— 1:250,000 Geological Series. Explan. Notes Bur. Miner. Resour. Geol. Geophys. Aust., SE/55-6. GOWER, J. C., 1966: Some distance properties of latent root and vector methods used in multivariate analysis. Biometrika, 53, pp. 325-38.

NUMERICAL ANALYSIS OF MODERN POLLEN SPECTRA KERSHAW, A. P., 1970: A pollen diagram from Lake Euramoo, north-east Queensland, Australia. New Phytol., 69, pp. 7 8 5 - 8 0 5 . , 1971: A pollen diagram from Quincan Crater, north-east Queensland, Australia. New Phytol., 70, pp. 669-81. LANCE, G . N . , & WILLIAMS, W . T., 1967: M i x e d -

data classificatory programs I. Agglomerative systems. Aust. Comput. J., 1, pp. 15-20. , , 1968: Note of new informationstatistic classificatory program. Comput. J., 11, p. 195. J., 1 9 5 9 : A physiognomic classification of Australian rain forests. J. Ecol., 47, pp.

WEBB, L .

551-70.

, 1968: Environmental relationships of the structural types of Australian rain-forest vegetation. Ecology, 49, pp. 296-311. A. P. Kershaw, Department of Biogeography & Geomorphology, Research School of Pacific Studies, Australian National University, P.O. Box 4, Canberra, A.C.T. 2600.

Spec.Publs geol.Soc.Aust., 4: pp. 191-199, 1973.

199

, TRACEY, J . G . , WILLIAMS, W . T . , & LANCE,

G. N., 1967a: Studies in the numerical analysis of complex rain-forest communities. I. A comparison of methods applicable to site/ species data. J. Ecol., 55, pp. 171-91. —, , , , 19676: Studies in the numerical analysis of complex rainforest communities. II. The problem of species-sampling. J. Ecol., 55, pp. 525-38. , , , , 1970: Studies in the numerical analysis of complex rain-forest communities. V. A comparison of floristic and physiognomic-structural data. J. Ecol., 58, pp. 203-232. WILLIAMS,

W.

T.,

LAMBERT,

J.

M.,

&

LANCE,

G. N., 1966: Multivariate methods in plant ecology. Similarity analyses and informationanalysis. J. Ecol., 54, pp. 427-46. WILLIAMS, W . T., & LANCE, G . N., 1968: Choice of strategy in the analysis of complex data. The Statistician, 18, pp. 31-44.

FURTHER DEVELOPMENTS IN COMPUTER ASSISTANCE TO POLLEN IDENTIFICATION By J. GUPPY, P. MILNE, M. GLIKSON, & H. MOORE (With 2 Text-Figures) ABSTRACT

The present paper describes recent changes and developments in a computerassisted storage and retrieval system for pollen morphological data (Walker et al., 1968) and illustrates the use of the substantial bank of stored reference data. A key for saccate grains and a sieving programme which retrieves from the deck all taxa bearing one or more specified characters are outlined. Selective coding of unknown grains which facilitates the reading of printed output and some preQuaternary interrogations and application are discussed. INTRODUCTION cise, and the diagrams in the key are wholly Quaternary palynology was developed in explanatory. The sections of the coding key Europe and North America and has there are listed below, followed by some characters, reached so sophisticated a level that there re- their assigned numbers and sections alongside main few outstanding difficulties in the produc- examples of grains exhibiting these characters. In the coding of reference pollen for the tion of basic pollen diagrams. In tropical areas and much of the southern hemisphere the flora store emphasis is placed on not losing information. If, for instance, both rounded and pointed is much less well known, often very different from the traditional areas of research, and colpi appear in the grains on a reference slide both these characters are recorded, in this often extremely complex. categories 128 and 129. Walker et al. (1968) described a computer pro- caseAsbya punching result of an interrogation the number gramme designed to ease the load on workers of sections in which at least one match between attempting identification of grains in these new categories is achieved determines the indiand complex situations, and here we draw viduals suggested and the order in which they attention to the changes and developments that are printed. The process is a 'forgiving' one; have been made since that publication, illus- that is, unlike a dichotomous key, a mistake trate the use we are making of the stored data, in observation or coding does not prevent the and outline proposals for further developments. chances of a match in any one section of the The system remains less comprehensive but coding key other characters are recorded in simpler and perhaps more practicable in use that section.if Sections A and B have an addithan that proposed by Germeraad & Muller tional directive function but even so if there (1970). is any doubt whether a grain is, say, tricolpate Basically the process consists of describing tricolporate both are recorded and both an 'unknown' pollen grain in a predetermined or these characters have a chance of being code and putting the coded description through in any interrogation. the computer to be compared with the coded matched The output has been organised so that it may descriptions of the modern reference pollen easily understood and worked from directly. collection of the Department of Biogeography be 'Rules of thumb' developed from use of the and Geomorphology of the Australian National over a period of time have been University. This process is termed an interro- programme incorporated to regulate the amount of output gation. from any one interrogation; it is important Figure 1 illustrates one page of the coding to print neither too much nor too little. key which consists of 149 characters divided into 17 sections. The method for measuring MODIFICATIONS TO THE the index polaris is shown in Figure 1 in Sec- PROGRAMME tion K. The coding procedure is not interpreThe paper by Walker et al. (1968) anticitative, the definitions of the characters are pre- pated the need to revise the key and print rules Spec.Publs geol.Soc.Aust., 4: pp. 201-206, 1973.

202

J. GUPPY, P. MILNE, M. GLIKSON, & H. M O O R E

12 SECTION H

EXINE ELEMENTS IN OPTICAL SECTION" (x 1600)

1 jx

71

Scabrate

72

Gemmate

)

73

Clavate

i

74

Verrucate ;

75

Baculate

{

76

Echinate

J

77

Psilate

(smoo

SECTION I

78

SCULPTURE ELEMENT DISTRIBUTION (excepting aperture areas) (x 1600)

U n i f o m all over

79

Size changes towards poles

80

Units change towards poles

81

Size changes towards apertures

82

Units change towards apertures

83

Size or units vary otherwise

SECTION J 84

+1

BXINE STRUCTURE

(x 1600)

layer

85

Clearly more than 1 layer

86

Uniform thickness

^ ^ ^

87

Thicker towards poles

88

Thinner towards poles

89

Clear columellae within the exine of the grain

SECTION K

INDEX POLARIS

(x 1600)

90

0

91

0 - .24

the circumference of which

92

.25- .49

unites the ends of the aperture

93

>

areas.

x is the diameter of a circle

.5

Measure 4 grains

x Lt

=

Index polaris

Fig. 1. A copy of one page of the coding key.

COMPUTER ASSISTANCE TO POLLEN IDENTIFICATION after a trial period. The number of sections has been increased from 11 to 17 and many characters in the original key are now defined more precisely. The increased number of sections has eliminated some obscurities which occurred with the use of the original key. As an example, one of the 11 sections contained characters for both the meridional colpus and the transverse aperture. It was thus impossible to determine to which aperture matches recorded within that section applied. In the present key this difficulty cannot occur. The flow chart remains essentially the same but print-out requirements are constantly being revised as special problems are encountered. For instance, as the number of reference grains coded in the data bank increased, the output was frequently cluttered by the printing of several closely similar species of a single genus matching in the same number of sections. This resulted in the loss of potentially valuable suggestions with only slightly lower match

203 scores and has been overcome by grouping such related individuals against the same serial number. This is illustrated in Figure 2 where pairs of such close (morphological) relatives are bracketed and together contribute only one to the total number sought. The same figure also demonstrates how, once a particular level of match score has been reached (in this case 11), the stop rule (20 units) is overridden and printing continues until all units with that score have been recorded.

It is also possible to obtain a list of reference individuals possessing or lacking one or more characters in common. For instance, all reference taxa with distorted colpi, other than those which also bear transverse apertures, may be retrieved by this sieving process. An additional subroutine entered from Section A has now been designed for use with saccate grains; the new key uses 56 characters divided into 11 sections.

Sections of Coding Key A. MAIN CLASS J. EXINE STRUCTURE B. APERTURE NUMBER AND K. INDEX POLARIS ARRANGEMENT L. MEASUREMENTS OF EXINE C. SHAPE IN LATERAL VIEW M. TRANSVERSE COLPUS OR PORUS D. SHAPE IN POLAR VIEW N. COLPUS STRUCTURE IN OPTICAL E. LAGUS SECTION F. LATUS O. COLPUS STRUCTURE IN TOP OR G. DEFINITIVE PATTERN IN TOP VIEW SURFACE VIEW H. EXINE ELEMENTS IN OPTICAL P. PORE STRUCTURE IN OPTICAL SECTION SECTION I. SCULPTURE ELEMENT DISTRIBUTION Q. PORE STRUCTURE IN TOP OR SURFACE VIEW POLLEN TAXA

SECTION

Eriocaulon hookerianum Artemisia

B B D

vulgaris

K K Boronia algida

M Pittosporum pullifolium Casuarinidites cainozoicus Casuarina sp. Triorites harrisii Asterolasia correifolia Dacrydium laxifolium

N

P J Saccate key

41

Spec.Publs geol.Soc.Aust., 4: pp. 201-206, 1973.

CHARACTERS — 34. Syncolpate: no 'island' at pole — 21. Tricolporate — 44. Inter semi-angular—apertures between corners — 92. Index polaris 13/45 = < 0.3 — 93. Index polaris 25/45 = > 0.5 (92 & 93 illustrate that relative length of colpus is indicated by recording index polaris) — 107. Transverse colpus (aperture length is ^ 2 x aperture width) — 119. Exine thicker adjacent to aperture.

140. Exine internally thickened in area approaching but not adjacent to aperture, but not forming a vestibulum. — 85. Clearly more than one layer. 39. Exine of cappa ± one-layered.

N> 2

(02A24001)

UNKNOWN INDIVIDUAL NO.

(TIME

171544,

DATE

02/04/71),

Ulmocese

10/29/35/42/56/62/67/77/78/84/86/93/95/100/103/139/142. (A 10) (Q

(B 2 9 )

(C 35)

(D 42)

(E 56)

(P 62)

(G 67)

(H 77)

(i 78)

MATCHES

IDENTIFICATION

~~3 4 5 6 7 8

13 13 12 12 12 12 12 11

1-0-0275-09-001-02 1-0-0060-04-001-08' 1-0-0045-01-009-08 1-0-0077-01-004-07 1-0-0155-14-001-07 1-0-0058-01-006-07 1-0-0058-04-001-07 1-0-0045-01-010-08

~

Ql

11

1-0-0058-02-002-08

9V 10 11 12 137 13( 14 15 16) 161 17

11 11 11 11 11 11 11 11 11 11 11

1-0-0058-02-004-07 1-0-0228-03-003-07 1-0-0060-01-003-08 1-0-0129-22-001-08 1-0-0249-04-001-08 1-0-0249-04-002-07 1-0-0249-02-002-07 1-0-0062-07-001-08 1-0-0149-09-002-08 1-0-0149-09-011-07 1-0-0055-04-001-07

18

11

1-0-0168-29-001-08

19 20 @

11 11 11

1-0-0062-12-001-07 1-0-0062-03-004-07 1-0-0052-01-003-07

Ftruth of th€ ti

good A 10 A 10 A 10 A 10 A 10 A 10 A 10 A 10 A 10 10 10 10 10 10 10 10 10 10 A 10 A 10 A 10 A 10 A 10 A 10

natch

B 10 C 10 B 10 C 10 01 B 10 10 B 10 10 B 10 10 B 10 01 B 10 01 B 10 01 B 10 01 B 10 10 B 10 01 B 10 01 B 10 10 B 10 01 B 10 01 B 10 10 B 10 01 B 10 10 B 10 01 B 10 01 B 10 10 B 10 01 B 10 01 B 10

(K 93)

Rphanarrtht (L 95 100 103)

/

port

\

O CJ

ANALYSIS OP MATCHES BY SECTION D 10 D 10 D 10 D 10 D 10 D 10 D 10 D 01 D 10 D 01 10 10 10 10 10 10 10 D 10 D 01 D 01 D 01 D 10 D 10 D 01

10 10 10 10 10 10 10 10 10 10 10 10 10 10 01 10 01 10 10 01 01 10

P 10 G 10 1 10 G 10 P 10 G 10 P 10 G 10 P 10 G 10 P 01 G 10 P 10 P 10 P 10 P 10 P 10 P 10 P 10 P 10 10 10 01 10 10 P 10 P 10 P 10 P 10 P 10

H 10 H 01 H 10 H 10 H 10 H 10 H 10 H 10 H 01 H 10 H 01 H 01 H 01 H 10 H 10 H 01 H 01 H 01 H 10 H 10 H 10 H 01 H 10 H 10

I 10 I 10 01 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10

J 20 J 20 J 20 J 20 J 20 J 20 20 20 20 20 11 20 20 20 20 20 11 20 20 20 20 20 20

matehts Fig. 2.

(P 139)

Extne t/iicknecs

142)

NO.

(J 84 86)

The result of an interrogation to the computer.

K 10 L 30 K 10 L 30 K 10 L 30 K 01 L 03 21 K 10 K 10 30 K 10 21 K 10 30 K 10 30 K 10 21 10 21 10 30 10 12 12 01 30 10 K 10 30 21 K 10 K 10 30 21 K 10 21 K 10 30 K 10 12 K 10 21 K 10 21 K 10

M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00 M 00

N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00 N 00

0 00 0 00 0 00 0 00 0 00 0 00 0 00 0 00 0 00 0 00 0 00 00 00 00 00 00 00 00 00 00 00 00 00 00

Q 10

P 10 10 10 01 01 01 01 01 01 01 01 10 01 01 01 10 01 01 01 01 10 01 01

Q 10 Q 10 Q 10 Q 01 Q 10 Q 10 Q 10 Q 10 Q 10 Q 10 Q 10 Q 10 Q 10 Q 01 Q 10 Q 10 Q 10 Q

10

Q Q Q Q Q

10

10 10 10 10

3

5 ffl

G r HH * C/3 O 2

s o o w

COMPUTER ASSISTANCE TO POLLEN IDENTIFICATION 205 VALIDATION OF THE CODING KEY significant so that priority was given to successThe reference bank now numbers 2500 ful matching in these sections when selecting pollen taxa. In 1970, when this number was a most likely reference individual from just over 2000, a check was made in our amongst those with highest sectional match laboratory to test the accuracy of our coding scores. This led to the microscopic examination and the specificity of the definitions in the key. of a slide of catalogue number 0060-4-1 Six members of the laboratory staff were in- (Ulmacea, Aphananthe sp.) underlined in volved, two of whom had done no coding Figure 2 which confirmed the convincing simibefore, and all worked only from the manual larity between it and the fossil grain. and key; no explanations were made available In another part of the description of the from more experienced colleagues. 'unknown' the code itself was used selectively. Twenty-five taxa from the reference bank The circularity of the pore (Section Q) was were selected for recoding. Each was coded by very clearly distinct and unusually symmetrical; three persons, one of whom in each case had no matter what other pore features in this secoriginally coded the grain for the reference tion {e.g. edges diffuse here and there) had bank. The aim was to check for uniformity matched with the reference descriptions (so among observers, and for changes in the per- ensuring a match in Section Q) a match with formance of a single observer over a period this circularity was essential to an acceptable identification. Accordingly no other characters of time. The coded descriptions were used to interro- within this section were recorded so ensuring gate the computer. Sixty-two of the 75 {i.e. that any match scored in this section was due 80 per cent) of the interrogations correctly re- to the circularity alone. called the original descriptions and at least some of the failures related to unusual mor- PRE-QUATERNARY APPLICATION phological features. In 22 cases the original Palynologists are sometimes unable to agree observers described the grains as they had done previously. This result gives considerable on the relative systematic positions of form confidence in the descriptive criteria currently taxa and their relation to the pollen grains of living plants. This kind of problem is exempliin use. fied by the reluctance to accept the affinity of Triorites harrisii with the grain of similar type, SELECTIVE CODING Casuarinidites cainozoicus. Preparations conA degree of flexibility in interrogation can taining both kinds from early and late Tertiary be achieved at either or both of two points, deposits of New South Wales (supplied by namely in the interpretation of the print-out Dr Helene Martin) were examined and several or by the selective use of the code itself. grains of each type coded. These descriptions Figure 2 may be used to illustrate both op- were then used to interrogate the modern tions; it is the printed output from an interro- reference pollen bank. The results led to the gation using a stephanoporate fossil grain from selection of the reference grain 45-1-1, Sirunki, New Guinea, the coded description of Casuarina sp., as the best match in each case which is printed at the top, followed, in the from a printed output in which, amongst others, line below, by the same code figures placed a number of Casuarina species appeared. The within the sections of the key. A list of sug- character 140 in Section P which led to selecgested identifications follows. Thus, serial num- tive coding of the 'unknowns' is defined in the ber 1 identification matched the 'unknown' in list printed earlier in the text. 13 sections, the appropriate reference indiviIt is therefore reasonable to conclude that dual is catalogued as 275-9-1 and the number Casuarinidites cainozoicus and Triorites harriof character matches obtained in each Section, sii, although differing from each other in detail, A-Q, is shown. In Section A, 10 indicates that are more like the same modern reference the 'unknown' grain is matched by the refer- pollen individual in our collection than either ence grain in one (1) character and unmatched of them is to any other, even to others of the in none (0); in Section J, 11 shows that the same genus {i.e. Casuarina). 'unknown' matched in one (1) character and Our experience with modern and Quaternary was unmatched in one (1). pollen leads us to suppose that a similar apHowever, in this particular instance, exine proach, indeed many of the same descriptive structure (Section J) and pore structure in categories, could profitably be used in the desoptical section (Section P) were considered cription and comparison of totally fossil palySpec.Publs geol.Soc.Aust., 4: pp. 201-206, 1973.

206

J. GUPPY, P. MILNE, M. GLIKSON, & H. MOORE

nomorphs. This has been confirmed by the successful coding of a number of early Tertiary grains (supplied by Mr I. Raine) mounted in a solid medium, and their use to interrogate the bank of reference descriptions. It does at least indicate not only the desirability but the feasibility of the kind of development envisaged as a second stage in the computerisation of palynology by Kremp (1970). CONCLUSIONS The system modified as described in this

paper continues to prove its value as an aid to the identification of Quaternary pollen grains. It seems to have potential uses, in its present or modified form in pre-Quaternary palynology and higher plant systematics. ACKNOWLEDGMENT We are indebted to Professor D. Walker and our colleagues in the Department for co-operation at various stages in the work.

REFERENCES Research Report No. 1, Department of Geosciences, University of Arizona, Tucson. puter based numerical coding system for the description of pollen and spores. Rev. Palaeo- WALKER, D . , MILNE, P., GUPPY, JOAN, & bot. Palyn., 10, pp. 175-202. WILLIAMS, JUDITH, 1968: The computer assisted storage and retrieval of pollen morKREMP, GERHARD O. W., 1970: Towards compuphological data. Pollen Spores, 10, pp. 251terisation of palynology-paleobotany: a pro62. gress report on a fact finding trip. Interim GERMERAAD, J. H., & MULLER, J., 1970: A c o m -

J. Guppy, M. Glikson, H. Moore, Department of Biogeography and Geomorphology, Research School of Pacific Studies, The Australian National University, P.O. Box 4, Canberra City, A.C.T. 2600.

P. Milne, Division of Computing Research, C.S.I.R.O., Canberra, A.C.T. 2600.

INDEX TO DESCRIBED AND/OR ILLUSTRATED SPECIES Page numbers in Italics refer to pages with citations of specimens in legends of plates or text-figures. Adenanthos barbigera 78 Adiantites lindsayoides 123 Aglaoreidia sp. A 72 Amanda fioribunda 124 Apiculatisporis bulliensis 154 Aratrisporites coryliseminis 155 A. parvispinosus 155 A. tenuispinosus 155 Asteropollis asteroides 17-18, 26, 32 Aulax pinifolia 78 Australopollis obscurus 21, 26, 52, 71 Avicennia-typz pollen 86 Barringtonia calyptrata 86 B. cf. calyptrata 86 B. sp. 86 Beauprea elegans 78 Beaupreaidites elegansiformis

72, 73-75

Cadargasporites senectus 154 C. sp. nov. 154 Camarozonosporites ohaiensis 71 C. sp. A 71 Canningia colliveri 118 Carnarvonia araliaefolia 78 Caryophyllidites polyoratus 71 Cicatricosisporites australiensis 118 C. hughesi 118 C. ludbrooki 118 Classopollis chateaunovi 129-130, 136, 139, 140 C. cf. chateaunovi 130, 139 C. meyeriana 130-132, 136, 139, 140 C. simplex 132, 136, 140 Clavatipollenites sp. 11, 26, 33 Cleistosphaeridium tenuum 164, 166 Coniopteris cf. hymenophylloides 121, 123 Convolutispora sp. 154 Cooksonites variabilis 118 Couperisporites tabulatus 118 Crustaesporites sp. 155 Crybelosporites stylosus 118 Cubiculosphaera maslinensis 164, 166 Cupanieidites orthoteichus 72 Cupuliferoidaepollenites cf. parvulus 12, 26, 33 Cyatheacidites annulata 51, 54 Cyathidites breviradiatus 154 Cycadopites follicularis 155 Cyclosporites hughesi 118 Cymatiosphaera? sp. 155 Deflandrea bakerii 169-170, 180, 182, 187 D. conorata 171-172, 179, 187 D. dartmooria 176, 180, 188 D. delineata 174, 180, 182, 188 D. cf. diebelii 179-180 D. dilwynensis 172, 179, 182, 187 D. druggii 171, 179, 187

D. extensa 178-179, 188 D. flounderensis 175-175, 779, 180, 188 D. heterophlycta 178, 182, 7). leptodermata 177, 752, 755 £>. medcalfii 175-176, 77P, 752, 755 D. obliquipes 173-174, 755 ZX pachyceros 173, 77P, 750, 752, 755 7>. pellucida 170-171, 757 D. pentaradiata 172-173, 779, 750, 752, 755 £>. phosphoritica 177-178, 750, 755 £>. robusta 173, 755 Z>. truncata 176-177, 779, 752, 755 Densoisporites playfordi 154 D. velatus 118 D. sp. nov. 154 Dictyotosporites speciosus 118 Dingodinium cerviculum 118 Discisporites psilatus 133, 136, 140 D. cf. psilatus 133, 136, 140 D. verrucosus 154 Drimys lanceolata 54 D. sp. 51, 54 Duplexisporites problematicus 154 Falcisporites australis 155 Fern-like foliage sp. 'b' 124 Foraminisporia asymmetricus 118 F. wonthaggiensis 118 Franklandia fucifolia 78 Geranium spp. 54 Grebespora concentrica 134, 136, 139 Grevillea punicea 78 'Guthoerlisporites' cancellosus 155 Hepaticites

discoides

120

Indospora clara 154 Ischyosporites sp. A 72 Januasporites spinulosus

118

Kraeuselisporites cuspidus 154 K. rallus 154 Kuylisporites waterbolkii 72 Laevigatosporites sp. 775 Liliacidites cf. intermedius 10-11, 26, 32 L. cf. kaitangataensis 9-10, 26, 32 L. sp. A 72 Lophotriletes novicus 154 Lueckisporites nyakapendensis 155 Lunatisporites noviaulensis 155 L. pellucidus 155 Lygistepollenites balmei 64, 77 L. ellipticus 64 L. florinii 64

208 Macadamia ternifolia 78 Malvacipollis diversus 72 Micrantheum hexandrum 54 M. sp. 52, 54 Micrhystridium? sp. 155 Monosulcites sp. A 72 Montia fontana 51, 54 Morkallacysta pyrctmidalis 163, 166 Muderongia tetracantha 118 Murospora florida 118 Myrtaceidites tenuis 72 'Nevesisporites' fossulatus 154 'N.' limatulus 154 N. vallatus 118 N. sp. nov. 154 N. sp. Nothofagidites asperus 64, 72 N. deminutus 66, 72 N. endurus 66, 71 N. falcatus 66, 72 N. flemingii 65-66, 71 N. goniatus 64-65, 72 N. senectus 71 Nypa-typc pollen 86 Nyssapollenites lanosus 19, 26, 34 N. squamosus 18-19, 26, 34 Odontochitina operculata 118 Ornamentifera sentosa 71 Pelargonium australe 54 P. rodneyanum 54 Peromonolites densus 71 Phimopollenites augathallaensis 17, 26, 33 P. pannosus 16-17, 26, 33 Phyllocladidites verrucosus 63, 71 Pilosisporites notensis 118 Polycingulatisporites sp. nov. 154 Polycolpites sp. A 71 Polypodiaceosisporites sp. nov. 154 Polypodiisporites mutabilis 154 Pometia pinnata 78 Protea tenax 78 Proteacidites annularis 71 P. asperopolus 66-67, 72 P. dilwynensis 72 P. grandis 72 P. hakeoides 75-76 P. pachypolus 72 P. palisadus 71 P. sp. A 71 P. sp. B 72 P. sp. 78 Protohaploxypinus cf. jacobii 155

INDEX P. microcorpus 155 P. reticulatus 155 P. samoilovichii 155 Quadrisporites horridus Quintinia sp. 52, 54

155

Reticuloidosporites arcus 118 Retusotriletes radiatus 154 Rewanispora foveolata 154 Rhizophoraceae-type pollen 86 Rousea georgensis 14, 26, 33 Rugulatisporites sp. A 72 Saeptodinium gravattensis 162, 166 S. tasmaniensis 163, 166 Santalumidites cainozoicus 72 Senectotetradites fistulosus 22, 26, 34 S. varireticulatus 21-22, 26, 34 Sonneratia-type pollen 86 Spinizonocolpites prominatus 67, 72 Stereisporites regium 71 S. (Tripunctisporis) sp. 71 Striatopollis cf. paraneus 15, 26, 33 Striomonosaccites morondavensis 155 Telopea speciosissima 78 Tetracolporites sp. A 71 Tigrisporites playfordi 154 Tiliaepollenites notabilis 72 Tricolpites cooksonae 13-14, 26, 34 T. geranioides 52-53. 54 T. gillii 71 T. longus 67, 71 T. minutus 12-13, 26, 33 T. waiparaensis 71 T. sp. A 71 T. sp. B 72 T. sp. 14, 26, 33 Tricolporites lilliei 67, 71 T. sp. A 72 Tricolporopollenites pelargonioides 53, 54 Trilobosporites purverulentus 118 Triorites edwardsii 71 T. harrisii 71 T. magnificus 72 T. minor 20, 26, 34 T. punctulatus 20, 26, 34 T. sp. 20-21, 26, 34 Triporopollenites sp. A 71 Triquitrites microgram]er 154 Verrucosisporites carnarvonensis Veryhachium? sp. 155 Vitreisporites pallidus 155 Welwitschiapites sp. nov.

155

154

AUTHOR INDEX

Citations in Italics refer to pages on which the references are set out. Authors may be cited more than once on the pages listed. Albrecht, L., 198 Croft, W. N., iv Amerada Petroleum Corporation of Australia Cronquist, A., 27, 30 Ltd, 115, 106 Crook, K. A. W., 49, 49 Archangelski, S., 129, 138 Cundill, Meyers & Associates, 89, 92, 106, 115 Auer, V., 81, 84 Australian Oil and Gas Corporation Ltd, 108 Dale, B., 159, 163, 165 Dale, M. B., 193, 198 Baker, G., iii, ix, 119, 125 P., 132, 138 Balme, B. E., ix, 5, 24, 29, 30, 83, 84, 87, 95, Danze-Corsin, Davey, R. J., 5, 30, 62, 70, 164, 177, 186 97, 98, 100, 102, 103, 106, 141, 143, 144, Davidson, S. E., 159, 165 146, 151, 151, 152 Davis, M. B., 191, 198 Banks, M. R., 143, 152 Day, R. W., 6, 7, 28, 30, 32, 89, 93, 100, 104, Barghoorn, E. S., 29, 31 106, 107, 108, 115 Barnard, P. D. W., 129, 138 Deane, H., 35, 50 Baur, G. N., 46, 47, 48, 49 Deflandre, G., ix, 70, 167, 168, 170, 171, 173, Beadle N. C. W., 46, 49 174, 178, 186 Belford, D. J., 100 de Jersey, N. J., xi, 87, 107, 127, 128, 130, 131, Benedek, S., 115, 107 132, 133, 134, 135, 137, 138, 144, 146, 151, Berggren, W. A., 61, 69 152, 155 Best, J. G., 192, 198 de Keyser, F., 192, 198 Boltenhagen, E., 25, 29 Denmead, A. K., 93, 107 Brenner, G. J., 11, 12, 13, 14, 15, 25, 27, 29 Dettmann, M. E., ix, xi, 3, 5, 7, 12, 16, 20, 21, Briche, P., 132, 138 23, 24, 28, 30, 56, 61, 70, 87, 93, 96, 97, 99, Brinkmann, R., 87, 106 101, 102, 103, 106, 106, 107, 119, 120, 122, Brown, B. R., 97, 107 125, 125, 128, 138 Brown, D. A., 49, 49 Doring, H., 70, 139 Bryan, R., 7, 31 Dorman, F. H., 28, 30, 48, 50 Burckle, L., 85 J. G., xi, 6, 7, 30, 119, 120, 122, 125, Burger, D., xi, 3, 5, 13, 15, 16, 17, 24, 25, 29, Douglas, 125, 170, 186 87, 92, 93, 97, 98, 106, 106, 115 Dow, D. B., 89, 106 Busnardo, R., 87, 106 Downie, C., 162, 164 Doyle, J. A., 11, 12, 13, 15, 18, 22, 24, 25, Cahoon, E. J., 25, 29 26, 27, 29, 30 Campbell, K. S. W., 49, 49 Drugg, W. S., 61, 70, 171, 186 Casey, D. J., 6, 32, 89, 107, 108, 109 Duigan, S. L., viii, ix Casey, J. N., 89, 106 Dulhunty, J. A., 143, 146, 152 Chaloner, W. G., 11, 25, 26, 29, 129, 139 Chandler, M. E. J., 84, 85 Chapman, F., vii Edwards, A. B., 119, 125 Chlonova, A. F., 101, 106 Eisenack, A., ix, x, 5, 6, 30, 61, 62, 69, 101, 102, Churchill, D. M., xi, 79, 81, 82, 83, 84, 85, 159, 106, 107, 167, 168, 172, 174, 176, 177, 178, 164 185, 186 Clarke, R. F. A., 5, 30 Erdtman, G., v, viii, 9, 18, 21, 26, 28, 30, 76, 77, 81, 85 Clausen, J., 79, 85 Clifford, H. T., viii Evans, P. R., xi, 5, 6, 7, 21, 30, 31, 55, 56, 70, Cockbain, A. E., 83, 84, 85 87, 89, 93, 95, 96, 97, 98, 99, 101, 102, 103, Conolly, J. R., 85 107, 115, 119, 126, 141, 143, 146, 152 Cookson, I. C., vii, viii, ix, x, 3, 5, 6, 12, 20, Evitt, W. R., 159, 165, 181, 186 30, 35, 49, 50, 51, 61, 62, 63, 64, 65, 66, 67, 69, Ewing, M., 85 70, 73, 74, 77, 97, 101, 102, 106, 107, 119, Exon, N. F., 6, 32, 89, 93, 98, 105, 107 125, 125, 167, 168, 170, 171, 172, 173, 174, 176, 177, 178, 185, 186 Faegri, K., 9, 30, 79, 85 Cornelius, K. D., 132, 139 Fiser, J., 50, 51 Couper, R. A., viii, 9, 10, 11, 19, 20, 24, 27, 30, Florin, R., 120, 126 50, 60, 63, 64, 65, 70, 73, 75, 77 Frazer, L., 48, 50 Cranwell, L. M., x, 64, 70 Freytag, I. B., 6, 30, 89, 107

INDEX 210 Kern County Land Company, 108 Galloway, M. C., 6, 32, 89, 100, 707, 108, 709 Kershaw, A. P., xi, 191, 199 Gamerro, J. C., 129, 138 Kirkegaard, A. G., 89, 707 Gay, L. O., 97, 707 Klaus, W., 127, 130, 131, 137, 138 Geiger, M. E., 132, 138 Geological Survey of Queensland, 89, 107, 115 Kohler, E., 50, 52 Kremp, G. O. W., 206, 206 Germeraad, J. H., 15, 30, 201, 206 Krutzsch, W., 12, 15, 31, 162, 165 Gill, E. D., 48, 50 Glaessner, M. F., 97 Laing, A. C. M., 93, 707, 108, 115 Glikson, M., xi, 201 Lambert, J. M., 199 Gocht, H., 101, 707, 159, 165, 177, 186 Lance, G. N., 193, 197, 198, 199 Goczan, F., 131, 139 Lang, W. H., iv, vii, viii Godwin, H., 82, 55 Lange, R. T., 160, 165 Gower, J. C., 196, 198 Lanjouw, J., 129, 138 Graham, B. K., 92, 707, 115 Lantz, J., 97, 108 Grebe, H., 143, 152 Larson, D. A., 8, 31 Gregory, C. M., 89, 105, 707, 115 Laveine, J. P., 132, 138 Grimes, K. G., 89, 707 Leffingwell, H. A., 8, 31 Groot, C. R., 12, 15, 17, 18, 25, 50, 80, 85 Groot, J. J., 12, 15, 17, 18, 25, 30, 80, 81, 85 le Hegarat, G., 87, 106 Leopold, E. B., 29, 31 Guillard, R. R. L., 159, 165 Leschik, G., 133, 138 Guinet, P., 81, 85 Leslie, R. B., 7, 31 Guppy, J., xi, 201, 206 Levet-Carette, J., 132, 138, 139 Lewis, J. H., 108, 115 Hamilton, M., 134, 138, 144, 146, 151, 152 Lindsay, J. M., 160, 161, 165 Hare, R., & Associates, 707, 115 Ludbrook, N. H., 6, 31 Harland, R., 159, 165 Lunt, C. K., 60, 70 Harris, W. F., viii Harris, W. K., a:, xi, 20, 21, 30, 35, 50, 57, 61, 63, Magloire, L., 15, 25, 27, 31 70, 159, 160, 165, 168, 186 Magne, J., 87, 106 Hawkins, P. J., 7, 31 Maitland, A. G., 83, 85 Helal, A. H., 25, 30 Helby, R., xi, 141, 143, 144, 146, 148, 151, Manum, S., ix Martin, A. R. H., xi, 73 152, 153, 155 Martin, H. A., xi, 35, 37, 40, 50, 51 Hennelly, J. P. F., 143, 146, 152, 153 Mazenot, G., 87, 108 Heslop-Harrison, J., 26, 31 McAndrews, J. H., 159, 165 Hiesey, W. M., 85 McElroy, C. T., 143, 146, 153 Hill, D., 89, 93, 707, 132, 138 McGowran, B., 61, 70, 160, 165 Hind, M. C., 144, 153 Mclntyre, D. J., 70 Hodgson, E. A., 98, 144, 153 McLennan, E., vii Hopkins, B. M., 55, 70 McPhee, I., 100, 108, 115 Hopping, C. A., 15, 30, 132, 138 McWhae, K. M., 47, 50 Hou, D., 82, 85 Medus, J., 128, 139 Houston, B. R., 160, 165 Medwell, L. M., 119, 122, 126 Huber, G., 159, 165 Milligan, E. N., 6, 32, 89, 107 Hughes, N. F., ix, 29, 31 Milne, P., xi, 201, 206 Mines Administration Pty Ltd, 108, 115 Ingram, B. S., 102, 707 Moore, M., xi, 201 Ingram, J. A., 89, 108 Mott, W. D., & Associates, 98, 108, 115 Irving, E., 28, 31, 82, 85 Muller, J., 11, 15, 18, 20, 25, 27, 29, 30, 31, 67, Iversen, J., 9, 30 70, 80, 81, 85, 201, 206 James, E. A., 56, 60, 70 Nilsson, O., 50, 51 Jansonius, J., 127, 129, 134, 138, 139 Nipkow, F., 159, 165 Jardine, S., 15, 25, 27, 31 Norris, G., 12, 13, 15, 17, 18, 25, 30, 97, 108, Jauncey, W., 89, 707, 108, 109, 115 159, 165 Jelgersma, S., 82, 85 Jenkins, D. G., 61, 70 Partridge, A. D., 43, 50 Johnson, N. E. A., 89, 108 Paten, R. J., 87, 89, 106, 707, 127, 138 Patton, R. T., 35, 50 Keck, D. D., 85 Kemp, E. M., 11, 12, 13, 24, 25, 31, 101, 707 Pels, S., 37, 50 Pemberton, R. L., 108, 115 Kenley, P. R., 119, 126

INDEX Penny, J. S., 12, 25, 50 Pettitt, J. M., 129, 759 Pflug, H. D., 74, 75, 77, 127, 129, 139 Pierce, R. L., 8, 12, 31 Pike, K. M., viii, 39, 50, 69 Playford, G., 3, 5, 7, 9, 12, 15, 20, 21, 23, 25, 28, 31, 87, 93, 96, 97, 101, 102, 103, 106, 107, 119, 120, 725, 132, 755, 759 Pocock, S. A. J., 24, 57, 97, 99, 101, 705, 127, 129, 139 Potonie, R., 8, 12, 15, 18, 19, 57, 63, 64, 70, 74, 77 Poumot, C., 98, 100, 117 Power, P. E., 93, 108 Pritchard, P. W., 89, 106 Reiser, R. F., 89, 705, 127, 132, 759 Reynolds, M. A., 7, 57, 89, 93, 106, 108 Reyre, Y., 127, 128, 129, 130, 131, 132, 134, 136, 137, 759 Richards, K. A., 55, 70 Richards, P. W., 82, 85 Ridley, H. N., 82, 85 Rouse, G. E., 15, 57 Ryan, J. C., 705, 115 Samoilovich, S. R., 73, 75, 77, 101 Sarjeant, W. A. S., 159, 162, 164, 765 Schulz, E., 24, 57, 132, 759 Scott, R. A., 29, 57 Senior, B. R., 89, 106, 707, 705, 115 Senior, D., 89, 705, 115 Seward, A. C., 119, 122, 125, 126 Singh, C., 12, 13, 14, 15, 17, 18, 25, 57, 101, 705 Singleton, O. P., viii, 119, 726, 168, 186 Skwarko, S. K., 5, 57 Smart, J., 89, 707 Smith, G. M., 160, 165 Srivastava, S. K., 8, 12, 14, 57, 73, 74, 75, 77 Staplin, F. L., 159, 165 Stover, L. E., xi, 5, 7, 21, 57, 55, 61, 70, 167 Strachan, I., 159, 165 Stuntz, J., 152 Takhatajan, A., 3, 21, 27, 29, 57 Taylor, D. J., 7, 28, 57, 56, 60, 70, 161, 165 Teichert, C., 96, 705 Te Punga, M. J., 70

211

Terpstra, G. R. J., 6, 57 Thiergart, F., 12, 18, 57 Thomas, B. M., 89, 705 Thomson, P. W., 12, 57, 74, 75, 77 Tracey, J. G., 799 Tralau, H., 122, 126 Traverse, A., 159, 165 Tschudy, B. D., 74, 75, 77 Turesson, G., 79, 55 Twist, R. F., 705, 115 Union Oil Development Corporation, 89, 705, 115 Valencia, M. J., 8, 57 van der Hammen, T., 8, 50 van Steenis, C. G. G. J., 79, 82, 55 Veevers, J. J., 97, 705 Venkatachala, B. S., 131, 759 Verdier, J. P., 5, 50 Vickery, J. W., 48, 50 Vine, R. R., 6, 52, 89, 100, 105, 706, 707, 705, 709, 115 Voisey, A. H., 143, 755 Walker, D., 50, 51, 193, 795, 201, 206 Wall, D., 159, 163, 765 Wallis, W. E., 55, 70 Wardle, P., 48, 50 Webb, L. J., 191, 192, 799 Weeks, L. G., 55, 70 Wells, A. T., 97, 705 White, M. E., 93, 709 Whitehouse, F. W., 87, 89, 709 Wijmstra, T. A., 8, 50 Williams, A. J., 127, 132, 759 Williams, G. L., 164 Williams, J., 206 Williams, W. T., 193, 197, 799 Williamson, W. H., 36, 50 Wilson, G. J., 61, 62, 70, 170, 756 Woods, J. T., 132, 755 Woolley, J. B., 89, 709 Wright, A. J., 152 Wright, C. W., 5, 6, 29, 32 Zaklinskaya, E. D., 25, 32 Zolnai, G., 98, 100, 709, 115, 117